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。     

Linking soil food web structure to above‐ and belowground ecosystem processes: a meta‐analysis

Soil fauna can be an important regulator of community parameters and ecosystem processes, but there have been few quantitative syntheses of the role of soil fauna in terrestrial soil communities and ecosystems. Here, we conducted a meta‐analysis to investigate the impacts of invertebrate soil micro‐ and mesofauna (grazers and predators) on plant productivity and microbial biomass. Overall our results indicate that an increase in the biomass of soil fauna increased aboveground plant productivity across ecosystems by 35% and decreased microbial biomass by 8%. In addition, we found no evidence for trophic cascades in terrestrial soil food webs, but the bacterivorous component of soil fauna influenced plant productivity and microbial biomass more than did the fungivorous component. Furthermore, changes in the biomass of soil fauna differentially affected plant productivity among plant functional groups: a higher biomass of soil fauna increased aboveground productivity by 70% in coniferous systems. However, in ecosystems dominated by legumes, a functional group with lower inorganic nitrogen requirements, there was no response of aboveground productivity to increases in the biomass of soil fauna. In sum, the results of this meta‐analysis indicate that soil fauna help to regulate ecosystem production, especially in nutrient‐limited ecosystems.     

Decomposition and nitrogen mineralization in natural and agroecosystems: the contribution of soil animals

The present article centres on the contribution of soil animals to organic. matter decomposition and nitrogen mineralization in natural and agro-ecosystems. Criteria are presented for the categorisation of the soil fauna in functional groups in order to be able to quantify the contribution of the soil fauna. Three types of classifications: size, habitat and food, are discussed. For various natural ecosystems, such as prairies and forests, and for agro-ecosystems a rather similar outcome of the faunal contribution to nitrogen mobilization of approximately 30% appears to exist. This value is dependent on various types of interactions among functional groups, changes in population density of microorganisms and soil fauna, seasonally changing abiotic factors and management, such as fertilization, harvesting and addition of harvest residues to the soil. Finally, to improve management of ecosystems as related to soil faunal activity in decomposition, lines are set out for further research such as the development of dynamic models, studies concerning the effects of perturbation in relation to microbial dominance and the integration of the study of below-ground food webs with ecological theories.     

陆地生态系统混合凋落物分解研究进展

凋落物分解在陆地生态系统养分循环与能量流动中具有重要作用,是碳、氮及其他重要矿质养分在生态系统生命组分间循环与平衡的核心生态过程。自然生态系统中,植物群落大多具有较高的物种丰富度和多样性,其混合凋落物在分解过程中也更有可能发生养分传递、化学抑制等种间互作,形成多样化的分解生境,多样性较高的分解者类群以及复杂的级联效应分解,这些因素和过程均对研究混合凋落物分解过程、揭示其内在机制形成了极大的挑战。从构成混合凋落物物种丰富度和多样性对分解生境、分解者多样性及其营养级联效应的影响等方面,综合阐述混合凋落物对陆地生态系统凋落物分解的影响,探讨生物多样性在凋落物分解中的作用。通过综述近些年的研究发现,有超过60%的混合凋落物对其分解速率的影响存在正向或负向的效应。养分含量有差异的凋落物混合分解过程中,分解者优先利用高质量凋落物,使低质量的凋落物反而具有了较高的养分有效性,引起低质量凋落物分解加快并最终使混合凋落物整体分解速率加快;而凋落物物种丰富度对土壤动物群落总多度有轻微的影响或几乎没有影响,但是对线虫和大型土壤动物的群落组成和多样性有显著影响,并随着分解阶段呈现一定动态变化;混合凋落物改变土壤微生物生存的理化环境,为微生物提供更多丰富的分解底物和养分,优化微生物种群数量和群落结构及其分泌酶的活性,并进一步促进了混合凋落物的分解。这些基于植物-土壤-分解者系统的动态分解过程的研究,表明混合凋落物分解作用不只是经由凋落物自身质量的改变,更会通过逐级影响分解者多样性水平而进一步改变分解速率和养分释放动态,说明生物多样性确实在一定程度上调控凋落物分解及其养分释放过程。     

Accumulation of local pathogens: a new hypothesis to explain exotic plant invasions

Recent studies have concluded that release from native soil pathogens may explain invasion of exotic plant species. However, release from soil enemies does not explain all plant invasions. The invasion of Ammophila arenaria (marram grass or European beach grass) in California provides an illustrative example for which the enemy release hypothesis has been refuted. To explore the possible role of plant–soil community interactions in this invasion, we developed a mathematical model. First, we analyzed the role of plant–soil community interactions in the succession of A. arenaria in its native range (north-western Europe). Then, we used our model to explore for California how alternative plant–soil community interactions may generate the same effect as if A. arenaria were released from soil enemies. This analysis was carried out by construction of a 'recovery plane' that discriminates between plant competition and plant–soil community interactions. Our model shows that in California, the accumulation of local pathogens by A. arenaria could result in exclusion of native plant species. Moreover, this mechanism could trigger the rate and spatial pattern of invasive spread generally observed in nature. We propose that our 'accumulation of local pathogens' hypothesis could serve as an alternative explanation for the enemy release hypothesis to be considered in further experimental studies on invasive plant species.     

白三叶(Trifolium repens)入侵对城市草坪生态系统土壤动物的影响

土壤动物是土壤生态系统中十分活跃的生物类群之一,也是城市草坪生态系统的重要组成部分。由于外来植物白三叶入侵,草坪土壤动物可能受到直接或间接的影响,进而改变城市草坪生态系统功能及过程。研究白三叶入侵对城市草坪生态系统土壤动物的影响,可为进一步了解外来植物入侵机制及城市草坪的建植养护提供理论依据。采用野外调查的试验方法对4种不同入侵程度下城市草坪的土壤动物群落特征和土壤理化性质进行了研究。结果如下：4种不同入侵程度的草坪样地共捕获土壤动物30099只,隶属于19目43科,其中小杆科、线蚓科和跳虫科为优势类群;土壤动物个体数量和类群数量整体表现为中度入侵 > 轻度入侵 > 对照 > 重度入侵,符合中度干扰假说;土壤动物个体数量和类群数量的季节动态主要表现为夏秋较高,冬春较低。CCA分析显示,土壤理化因子中铵态N对土壤动物影响相对较小,可能与白三叶改善了草坪土壤的供N水平并使其不成为土壤动物分布的限制因子有关;同时,土壤理化因子对土壤动物优势类群影响较小,而主要影响常见和稀有类群。总体而言,一定程度的白三叶入侵增加了土壤动物群落的多样性,但当达到重度入侵时,白三叶形成单优群落,减少了植物多样性,进而使土壤动物可获得的生活资源减少,土壤动物群落多样性呈下降趋势;白三叶入侵后通过改变土壤理化性质,影响到土壤动物常见和稀有类群,并最终改变草坪土壤动物的群落结构。     

Interactions between mycorrhizal fungi and other soil organisms

Mycorrhizal fungi interact with a wide range of other soil organisms, in the root, in the rhizosphere and in the bulk soil. These interactions may be inhibitory or stimulatory; some are clearly competitive, others may be mutualistic. Effects can be seen at all stages of the mycorrhizal fungal life-cycle, from spore population dynamics (predation, dispersal and germination) through root colonization to external hyphal growth. Two areas that seem likely to be of particular importance to the functioning of the symbiosis are the role of bacteria in promoting mycorrhiza formation and of soil animals in grazing the external mycelium. Mycorrhizal fungi also modify the interactions of plants with other soil organisms, both pathogens, such as root-inhabiting nematodes and fungi, and mutualists, notably nitrogen-fixing bacteria. These interactions are probably important both in natural ecosystems, where pathogens are increasingly recognized as playing controlling roles, and in agricultural systems, where mycorrhizas may be valuable in designing integrated systems of pest control and growth stimulation.     

Managing the ecology of foliar pathogens: ecological tolerance in crops

The phyllosphere is a rich and varied microbial community comprising organisms with diverse functional types. Its composition is strongly influenced by both genotypic and environmental factors, many of which can be manipulated by breeding, agronomy and crop protection strategies in an agricultural context. These factors also affect the complex interactions between the microbes, which in turn affect their interaction with their host plant. Whether or not an organism becomes pathogenic and the subsequent expression of disease are also influenced by all these factors. Understanding the population dynamic balance between the organisms of the phyllosphere as an ecological system should lead to new approaches in agronomy, crop protection and breeding that enhance sustainability, where the previously presumed requirement to eliminate putative pathogens is replaced by management that favours dominance of beneficial organisms and contains putative pathogens in asymptomatic or stable states.     

Plants respond to pathogen infection by enhancing the antifungal gene expression of root-associated bacteria

Plant health and fitness widely depend on interactions with soil microorganisms. Some bacteria such as pseudomonads can inhibit pathogens by producing antibiotics, and controlling these bacteria could help improve plant fitness. In the present study, we tested whether plants induce changes in the antifungal activity of root-associated bacteria as a response to root pathogens. We grew barley plants in a split-root system with one side of the root system challenged by the pathogen Pythium ultimum and the other side inoculated with the biocontrol strain Pseudomonas fluorescens CHA0. We used reporter genes to follow the expression of ribosomal RNA indicative of the metabolic state and of the gene phlA, required for production of 2,4-diacetylphloroglucinol, a key component of antifungal activity. Infection increased the expression of the antifungal gene phlA. No contact with the pathogen was required, indicating that barley influenced gene expression by the bacteria in a systemic way. This effect relied on increased exudation of diffusible molecules increasing phlA expression, suggesting that communication with rhizosphere bacteria is part of the pathogen response of plants. Tripartite interactions among plants, pathogens, and bacteria appear as a novel determinant of plant response to root pathogens.     

Friends and foes: streptomycetes as modulators of plant disease and symbiosis

The ecological role of soil streptomycetes within the plant root environment is currently gaining increased attention. This review describes our recent advances in elucidating the complex interactions between streptomycetes, plants, pathogenic and symbiotic microorganisms. Streptomycetes play diverse roles in plant-associated microbial communities. Some act as biocontrol agents, inhibiting plant interactions with pathogenic organisms. Owing to the antagonistic properties of streptomycetes, they exert a selective pressure on soil microbes, which may not always be for plant benefit. Others promote the formation of symbioses between plant roots and microbes, and this is in part due to their direct positive influence on the symbiotic partner, expressed as, e.g., promotion of hyphal elongation of symbiotic fungi. Recently, streptomycetes have been identified as modulators of plant defence. By repressing plant responses to pathogens they facilitate root colonisation with pathogenic fungi. In contrast, other strains induce local and systemic resistance against pathogens or enhance plant growth. In conclusion, while streptomycetes have a clear potential of acting as biocontrol agents, care has to be taken to avoid strains that select for virulent pathogens or enhance disease development. We argue towards the use of an integrated screening approach in the search for efficient biocontrol agents, including assays on in vitro antagonism, plant growth, and disease suppression.     

高原鼢鼠干扰下高寒草甸大型土壤动物多样性对环境因子的响应

为研究高寒草甸大型土壤动物群落组成和分布对环境因子的响应,选取祁连山东段的甘肃省天祝县高原鼢鼠典型分布区域,以鼠丘密度代表干扰强度设置4个干扰区。调查各干扰区植被群落特征、土壤理化性质、大型土壤动物类群组成及其数量,采用约束性排序方法分析环境因子对大型土壤动物类群组成和分布的影响。结果表明：高原鼢鼠干扰下高寒草甸大型土壤动物优势类群为瓦娄蜗牛科、象甲科和短角亚目幼虫;极重度干扰区大型土壤动物类群的丰度、丰富度、Shannon指数显著高于重度干扰区（P<0.05）;多元回归分析表明大型土壤动物类群丰度、丰富度和Shannon指数与土壤温度呈显著负相关（P<0.05）,丰富度与土壤含水量呈显著负相关（P<0.05）,丰度与土壤紧实度呈显著负相关（P<0.05）,丰富度和Shannon指数与植物Shannon指数呈显著负相关（P<0.05）;冗余分析和偏冗余分析表明,土壤温度、紧实度和含水量是影响高寒草甸大型土壤动物类群组成和分布的主要环境因子。     

生物炭介导植物病害抗性及作用机理

生物炭是生物有机材料在缺氧或限氧条件下经高温热裂解后生成的固体产物,在固碳减排、污染修复、土壤改良等方面具有较大的应用潜力。研究表明,生物炭在植物病害胁迫中也起重要的抗性作用。综述了国内外关于生物炭缓解植物病害的相关研究,重点介绍了生物炭在降低病害和提高植物抗性方面的作用机理。生物炭通过诱导植物增强系统抗性,改良土壤理化特性,改变土壤微生物群落结构,增加土壤有益微生物类群的丰度和活性,吸附病原菌及其产生的有毒物质等来降低病原菌对寄主植物的侵害作用,从而促进植物生长和增强植株抗病性。生物炭对病害的抗病效果与生物炭的原料类型、用量、土壤及病害类型等有关。未来的研究应重点应围绕"生物炭-土壤-植物病害"体系,借助组学手段,深入研究生物炭介导植物病害的分子机理。     

锡林郭勒草原自然保护区土壤动物初步调查

土壤动物的生存、活动与栖息地环境密切相关。一方面栖息地生态因子决定着土壤动物的生存与活动;另一方面,土壤动物对土壤有机质的改变、土壤微生物的生存、植物的生长等诸生态因子的变化起着一定的促进作用。因此随着草原生态系统研究的日趋深入,土壤动     

Soil microbial communities along elevational gradients in the Madrean Sky Islands

The Madrean Sky Islands are mountain ranges isolated by a ‘desert sea’. This area is a biodiversity hotspot currently threatened by climate change. Here, we studied soil microbial communities along elevational gradients in eight Madrean Sky Islands in southeastern Arizona (USA). Our results showed that while elevational microbial richness gradients were weak and not consistent across different mountains, soil properties strongly influenced microbial community composition (overall composition and the abundance of key functional groups) along elevational gradients. In particular, warming is associated with a higher abundance of soil-borne fungal plant pathogens that concomitantly might facilitate upward elevational shifts of plant species released from negative plant–soil feedbacks. Furthermore, projected warming and drought in the area aggravated by anthropogenic nitrogen deposition on mountain tops (and thus, decreasing nitrogen limitation) can enhance a shift from ectomycorrhizal to arbuscular mycorrhizal fungi. Overall, these results indicate that climate change effects on plant–soil interactions might have profound ecosystem consequences.     

土壤动物多样性的地理分布及其生态功能研究进展

土壤动物多样性地理分布及其生态功能研究已成为地学和生态学等领域共同关注的科学前沿。本文在介绍相关研究最新进展的基础上, 讨论已有研究的局限性或不确定性, 展望未来研究的重点方向。近10年来, 代表性土壤动物类群的全球分布研究取得突破性进展; 国内土壤动物研究的尺度和采样区域也有明显拓展, 尤其在蚯蚓和线虫相关研究上取得了系列成果。结果表明, 土壤动物多样性随纬度的变化模式主要有两种, 即在低纬度的热带最高或在中纬度的温带最高; 而土壤动物多度与多样性可能同步变化、无明显关系、截然不同甚至相反; 降水、植物生产力和土壤有机质是土壤动物分布格局的关键驱动力, 但它们的影响力因土壤动物类群不同而异。土壤动物具有改善土壤物理结构、促进养分循环和有机碳稳定、提高作物健康水平等多重功能; 土壤动物的多功能性评估方兴未艾, 但仍面临诸多挑战。简单分析土壤动物随经纬度等的变化规律存在较大局限性, 考虑在基于地质-生态历史及“经纬度-海拔-离海岸距离”等构建的多维时空框架内, 探究土壤动物分布特征及其驱动力。土壤动物分布格局对其潜在的生态功能有关键影响, 但是目前对土壤动物分布格局的预测和模拟仍主要依靠经验模型; 代谢生态学等理论在土壤动物群落研究中的应用值得关注。探究分类多样性的冗余机制, 突出功能多样性, 可以将生物多样性与生态功能更好地联系起来; 同时, 需要在特定条件和时空下, 从整个土壤食物网及其与植物的联系中理解土壤动物多样性与多功能性的联系。建议未来关注两个研究方向: (1)量化人类活动和气候变化给土壤动物多样性和生态功能带来的巨大不确定性; (2)完善土壤动物群落特征预测的理论框架和开展土壤动物群落的精准调控, 综合评价其多功能性, 进而将土壤动物与人类福祉更紧密地联系起来。     

Effects of soil decomposer invertebrates (protozoa and earthworms) on an above-ground phytophagous insect (cereal aphid) mediated through changes in the host plant

We investigated if the activity of soil invertebrates (protozoa and earthworms) affected the performance of barley and if effects propagated higher up the above-ground food chain into herbivores (cereal aphid, Sitobion avenae ). Barley plants were grown individually in microcosms containing defaunated soil and grass residues. Plants were grown in soil containing: a) no added fauna, b) protozoa, c) earthworms, or d) protozoa and earthworms. After 7 weeks growth at 20°C three adult cereal aphids were added to each plant on separate leaves. The aphids were allowed to grow and reproduce for another 2 weeks before the experiment was destructively sampled. Amounts of mineral N in the soil and leached from the microcosms were significantly reduced by the presence of soil animals. Correspondingly plant biomass and total plant N content were increased significantly by soil animals, protozoa in particular. The different mechanisms responsible for changes in nutrient turnover in presence of protozoa and earthworms are discussed. Aphid performance was strongly influenced by the presence of protozoa, but not by earthworms. In the presence of protozoa the numbers and biomass of adult and juvenile aphids were significantly increased. These effects are likely due to an increased N content in barley plants and consequently increased nitrogen availability to aphids. The results underline that the detritivore and herbivore systems are intimately linked.     

No adverse effect of genetically modified antifungal wheat on decomposition dynamics and the soil fauna community--a field study

The cultivation of genetically modified (GM) plants has raised several environmental concerns. One of these concerns regards non-target soil fauna organisms, which play an important role in the decomposition of organic matter and hence are largely exposed to GM plant residues. Soil fauna may be directly affected by transgene products or indirectly by pleiotropic effects such as a modified plant metabolism. Thus, ecosystem services and functioning might be affected negatively. In a litterbag experiment in the field we analysed the decomposition process and the soil fauna community involved. Therefore, we used four experimental GM wheat varieties, two with a race-specific antifungal resistance against powdery mildew (Pm3b) and two with an unspecific antifungal resistance based on the expression of chitinase and glucanase. We compared them with two non-GM isolines and six conventional cereal varieties. To elucidate the mechanisms that cause differences in plant decomposition, structural plant components (i.e. C∶N ratio, lignin, cellulose, hemicellulose) were examined and soil properties, temperature and precipitation were monitored. The most frequent taxa extracted from decaying plant material were mites (Cryptostigmata, Gamasina and Uropodina), springtails (Isotomidae), annelids (Enchytraeidae) and Diptera (Cecidomyiidae larvae). Despite a single significant transgenic/month interaction for Cecidomyiidae larvae, which is probably random, we detected no impact of the GM wheat on the soil fauna community. However, soil fauna differences among conventional cereal varieties were more pronounced than between GM and non-GM wheat. While leaf residue decomposition in GM and non-GM wheat was similar, differences among conventional cereals were evident. Furthermore, sampling date and location were found to greatly influence soil fauna community and decomposition processes. The results give no indication of ecologically relevant adverse effects of antifungal GM wheat on the composition and the activity of the soil fauna community.     

Linking soil biodiversity and vegetation: implications for a changing planet

Soil biota are intimately tied to plant communities through herbivory and symbiosis and indirectly by the decomposition of dead organic plant material. Through both roots and aboveground organic material (e.g., leaves and wood), plants provide substantial inputs of organic matter to soil systems. Plants are the basis for most biotic soil food webs that comprise an enormous diversity of species whose multiple interactions function to help regulate nutrient cycling, which in turn influences plant growth. Many factors govern the biogeography of soil biota, including the physical and chemical properties of soil, climate, the composition and type of vegetation, and interactions with other soil biota. Despite awareness of factors influencing soil communities, no single factor allows predictions of soil animal diversity or distribution. However, research is showing that plants can have unique soil biotic communities. Degradation of soil, which removes predators and biotic regulation that occurs in less managed ecosystems, can result in increased pathogens and pests that affect humans, other animals and plants. Global changes such as land use, desertification, and soil pollution all have been shown to alter soil animal diversity and abundance. Because of our dependence on soils and plant production, studies linking soil biotic communities to primary productivity are needed to assure long-term soil sustainability.     

Responses of soil biota to elevated atmospheric carbon dioxide

Increasing concentrations of atmospheric CO₂ could have dramatic effects upon terrestrial ecosystems including changes in ecosystem structure, nutrient cycling rates, net primary production, C source-sink relationships and successional patterns. All of these potential changes will be constrained to some degree by below ground processes and mediated by responses of soil biota to indirect effects of CO₂ enrichment. A review of our current state of knowledge regarding responses of soil biota is presented, covering responses of mycorrhizae, N-fixing bacteria and actinomycetes, soil microbiota, plant pathogens, and soil fauna. Emphasis will be placed on consequences to biota of increasing C input through the rhizosphere and resulting feedbacks to above ground systems. Rising CO₂ may also result in altered nutrient concentrations of plant litter, potentially changing decomposition rates through indirect effects upon decomposer communities. Thus, this review will also cover current information on decomposition of litter produced at elevated CO₂. Summary Predictably, the responses of soil biota to CO₂ enrichment and the degree of experimental emphasis on them increase with proximity to, and intimacy with, roots. Symbiotic associations are all stimulated to some degree. Total plant mycorrhization increases with elevated CO₂. VAM fungi increase proportionately with fine root length/mass increase. ECM fungi, however, exhibit greater colonization per unit root length/mass at elevated CO₂ than at current atmospheric levels. Total N-fixation per plant increases in all species examined, although the mechanisms of increase, as well as the eventual benefit to the host relative to N uptake may vary. Microbial responses are unclear. The assumption that changes in root exudation will drive increased mineralization and facilitate nutrient uptake should be examined experimentally, in light of recent models. Microbial results to date suggest that metabolic activity (measured as changes in process rates) is stimulated by root C input, rather than population size (measured by cell or colony counts). Insufficient evidence exists to predict responses of either soil-borne plant pathogens or soil fauna (i.e., food web responses). These are areas requiring attention, the first for its potential to limit ecosystem production through disease and the second because of its importance to nutrient cycling processes. Preliminary data on foliar litter decomposition suggests that neither nutrient ratios nor decomposition rates will be affected by rising CO₂. This is another important area that may be better understood as the number of longer term studies with more realistic CO₂ exposures increase. Evidence continues to mount that C fixation increases with CO₂ enrichment and that the bulk of this C enters the belowground component of ecosystems. The global fate and effects of this additional C may affect all hierarchical levels, from organisms to ecosystems, and will be largely determined by responses of soil biota.     

土壤生物多样性对植物利用营养物质和水分的影响

土壤生物多样性影响土壤营养物质和水等自然资源的有效利用是农业生态系统的重要功能.有证据表明土壤微生物多样性可提高土壤营养物质和水的利用率.土壤动物对土壤营养物质和水的有效利用也产生了明显的影响,它主要是通过促进营养物质的转化和改善土壤水渗透率而间接实现的.从已有文献的报道来看,单从某一个方面来研究土壤生物多样性对土壤营养物质和水的有效利用的影响并不能全面地反应它们之间相互作用的规律.因此未来的研究应当把土壤生物多样性、土壤营养物质和水的供给与植物生物多样性、植物营养物质和水的利用率结合起来,这样才有可能从理论上对土壤生物多样性与营养物质和水的利用之间的关系有更加深刻的认识并运用到实践中.