Soil microbes and the availability of soil nutrients

It is likely to provide plants with their necessary nutrients using chemical and biological fertilization. Although chemical fertilization is a quick method, it is not recommendable economically and environmentally, especially if overused. Biological fertilization is the use of soil microbes including arbuscular mycorrhizal fungi and plant growth promoting rhizobacteria to inoculate plants. It has been proved that biological fertilization is an efficient method to supply plants with their necessary nutrients. It is economically and environmentally recommendable, because it results in sustainability. In this article, some of the most important details including the mechanisms and processes regarding the effects of soil microbes on the availability and hence uptake of nutrients by plant are reviewed. Such details can be important for the selection and hence production of microbial inoculums, which are appropriate for biological fertilization.     

土壤微生物在植物获得养分中的作用

大量施用化肥是当今农业的一个重要特征。化肥为粮食增产做出了巨大贡献,同时也带来一系列问题,如土壤酸化、水体富营养化、温室气体排放、资源耗竭等,直接威胁着农业可持续发展。土壤微生物是陆地生态系统植物多样性和生产力的重要驱动者,直接参与了植物获得养分和土壤养分循环两个过程。因此,通过调控土壤微生物的功能,有望降低农业对化肥的过分依赖。介绍了共生固氮菌、菌根真菌和根际促生菌对植物获得养分能力的影响及其机制,分析了土壤微生物对土壤氮、磷循环的影响及其与土壤养分生物有效性、养分损失的关系。依据这些知识,提出了改善植物营养、降低化肥施用的土壤微生物途径。虽然大量试验已证明了土壤微生物在改善植物营养中的重要作用,但是大面积应用土壤微生物技术来改善植物营养还存在不少问题。随着以后对这方面研究的加强以及上述问题的不断解决,土壤微生物有望在降低化肥施用量和维持农业可持续发展中做出重要贡献。     

植物与土壤微生物在调控生态系统养分循环中的作用

陆地生态系统的地上、地下是相互联系的。植物与土壤微生物作为陆地生态系统中的重要组成部分, 它们之间的相互作用是生态系统地上、地下结合的重要纽带。该文首先介绍了植物在养分循环中对营养元素的吸收、积累和归还等作用, 阐述了土壤微生物对养分有效性及土壤质量具有重要的作用。其次, 重点综述了植物与土壤微生物之间相互依存、相互竞争的关系。植物通过其凋落物与分泌物为土壤微生物提供营养, 土壤微生物作为分解者提供植物可吸收的营养元素, 比如共生体菌根真菌即可使植物根与土壤真菌达到互惠。然而, 植物的养分吸收与微生物的养分固持同时存在, 因而两者之间存在对养分的竞争。通过植物多样性对土壤微生物多样性的影响分析, 以及土壤微生物直接或间接作用于植物多样性和生产力的分析, 探讨了植物物种多样性与土壤微生物多样性之间的内在联系。针对当前植物与土壤微生物对养分循环的调控机制的争论, 提出植物凋落物是调节植物与土壤微生物养分循环的良好媒介, 植物与土壤微生物的共同作用对维持整个生态系统的稳定性具有重要意义。也指出了目前在陆地生态系统地上、地下研究中存在的不足和亟待解决的问题。     

Arbuscular mycorrhizal fungi and nitrogen uptake

Nitrogen (N) is among the most important macro-nutrients significantly affecting plant growth and yield production. Accordingly, N must be supplied adequately so that optimum amounts of yield are resulted. There are different ways of supplying N to the plant including the use of chemical and biological fertilization. The chemical properties of N make it very mobile, especially under humid conditions. Hence, N must not be overfertilized with respect to the economical and environmental points of view. N Biological fertilization includes the use of plant growth-promoting rhizobacteria (PGPR) including the N-fixing bacteria, rhizobium. There are also arbuscular mycorrhizal (AM) fungi in the soil, which are symbiotic to most terrestrial plants enhancing plant growth and yield production through increasing the uptake of water and nutrients by the host plant. Numerous experiments have indicated the important role of AM fungi in enhancing P uptake by plant. However, it is yet a matter of debate that how AM fungi may affect soil N dynamic and hence plant N uptake. Some of the most important and recent aspects regarding such effects by AM fungi are highlighted, which can be of significance to health and productivity of the ecosystem.     

Biological costs and benefits to plant-microbe interactions in the rhizosphere

This review looks briefly at plants and their rhizosphere microbes, the chemical communications that exist, and the biological processes they sustain. Primarily it is the loss of carbon compounds from roots that drives the development of enhanced microbial populations in the rhizosphere when compared with the bulk soil, or that sustains specific mycorrhizal or legume associations. The benefits to the plant from this carbon loss are discussed. Overall the general rhizosphere effect could help the plant by maintaining the recycling of nutrients, through the production of hormones, helping to provide resistance to microbial diseases and to aid tolerance to toxic compounds. When plants lack essential mineral elements such as P or N, symbiotic relationships can be beneficial and promote plant growth. However, this benefit may be lost in well-fertilized (agricultural) soils where nutrients are readily available to plants and symbionts reduce growth. Since these rhizosphere associations are commonplace and offer key benefits to plants, these interactions would appear to be essential to their overall success.     

Mechanisms of action of plant growth promoting bacteria

The idea of eliminating the use of fertilizers which are sometimes environmentally unsafe is slowly becoming a reality because of the emergence of microorganisms that can serve the same purpose or even do better. Depletion of soil nutrients through leaching into the waterways and causing contamination are some of the negative effects of these chemical fertilizers that prompted the need for suitable alternatives. This brings us to the idea of using microbes that can be developed for use as biological fertilizers (biofertilizers). They are environmentally friendly as they are natural living organisms. They increase crop yield and production and, in addition, in developing countries, they are less expensive compared to chemical fertilizers. These biofertilizers are typically called plant growth-promoting bacteria (PGPB). In addition to PGPB, some fungi have also been demonstrated to promote plant growth. Apart from improving crop yields, some biofertilizers also control various plant pathogens. The objective of worldwide sustainable agriculture is much more likely to be achieved through the widespread use of biofertilizers rather than chemically synthesized fertilizers. However, to realize this objective it is essential that the many mechanisms employed by PGPB first be thoroughly understood thereby allowing workers to fully harness the potentials of these microbes. The present state of our knowledge regarding the fundamental mechanisms employed by PGPB is discussed herein.     

The role of mycorrhizae and plant growth promoting rhizobacteria (PGPR) in improving crop productivity under stressful environments

Both biotic and abiotic stresses are major constrains to agricultural production. Under stress conditions, plant growth is affected by a number of factors such as hormonal and nutritional imbalance, ion toxicity, physiological disorders, susceptibility to diseases, etc. Plant growth under stress conditions may be enhanced by the application of microbial inoculation including plant growth promoting rhizobacteria (PGPR) and mycorrhizal fungi. These microbes can promote plant growth by regulating nutritional and hormonal balance, producing plant growth regulators, solubilizing nutrients and inducing resistance against plant pathogens. In addition to their interactions with plants, these microbes also show synergistic as well as antagonistic interactions with other microbes in the soil environment. These interactions may be vital for sustainable agriculture because they mainly depend on biological processes rather than on agrochemicals to maintain plant growth and development as well as proper soil health under stress conditions. A number of research articles can be deciphered from the literature, which shows the role of rhizobacteria and mycorrhizae alone and/or in combination in enhancing plant growth under stress conditions. However, in contrast, a few review papers are available which discuss the synergistic interactions between rhizobacteria and mycorrhizae for enhancing plant growth under normal (non-stress) or stressful environments. Biological interactions between PGPR and mycorrhizal fungi are believed to cause a cumulative effect on all rhizosphere components, and these interactions are also affected by environmental factors such as soil type, nutrition, moisture and temperature. The present review comprehensively discusses recent developments on the effectiveness of PGPR and mycorrhizal fungi for enhancing plant growth under stressful environments. The key mechanisms involved in plant stress tolerance and the effectiveness of microbial inoculation for enhancing plant growth under stress conditions have been discussed at length in this review. Growth promotion by single and dual inoculation of PGPR and mycorrhizal fungi under stress conditions have also been discussed and reviewed comprehensively.     

有机无机肥配施提高麦-稻轮作系统中水稻氮肥利用率的机制

采用盆栽试验,研究了有机无机肥配施对麦-稻轮作系统中水稻氮素累积动态和土壤氮素供应动态的影响,并从微生物学角度探讨了有机无机肥协同提高水稻氮肥利用率的机制.结果表明：有机无机肥配施处理的土壤微生物生物量碳、氮和矿质态氮在水稻分蘖期前低于化肥处理,而在抽穗期至灌浆期显著高于其他处理.土壤氮素供应动态与水稻吸收利用氮素规律吻合程度最高,促进了水稻产量、生物量和氮素累积量的增加,显著提高了水稻的氮肥利用率.其主要机制是有机无机肥配施促进了土壤微生物繁殖,使其在水稻生育前期固持了较多的矿质氮,在水稻生育中、后期这些氮素逐渐被释放以供水稻吸收利用,较好地满足了水稻各阶段生长发育对氮素养分的需求.     

Progressive N limitation of plant response to elevated CO2: a microbiological perspective

A major uncertainty in predicting long-term ecosystem C balance is whether stimulation of net primary production will be sustained in future atmospheric CO₂ scenarios. Immobilization of nutrients (N in particular) in plant biomass and soil organic matter (SOM) provides negative feedbacks to plant growth and may lead to progressive N limitation (PNL) of plant response to CO₂ enrichment. Soil microbes mediate N availability to plants by controlling litter decomposition and N transformations as well as dominating biological N fixation. CO₂-induced changes in C inputs, plant nutrient demand and water use efficiency often have interactive and contrasting effects on microbes and microbially mediated N processes. One critical question is whether CO₂-induced N accumulation in plant biomass and SOM will result in N limitation of microbes and subsequently cause them to obtain N from alternative sources or to alter the ecosystem N balance. We reviewed the experimental results that examined elevated CO₂ effects on microbial parameters, focusing on those published since 2000. These results in general show that increased C inputs dominate the CO₂ impact on microbes, microbial activities and their subsequent controls over ecosystem N dynamics, potentially enhancing microbial N acquisition and ecosystem N retention. We reason that microbial mediation of N availability for plants under future CO₂ scenarios will strongly depend on the initial ecosystem N status, and the nature and magnitude of external N inputs. Consequently, microbial processes that exert critical controls over long-term N availability for plants would be ecosystem-specific. The challenge remains to quantify CO₂-induced changes in these processes, and to extrapolate the results from short-term studies with step-up CO₂ increases to native ecosystems that are already experiencing gradual changes in the CO₂ concentration.     

根系代谢物介导的植物-微生物互作的研究进展

植物根系代谢物是植物-微生物互作的桥梁纽带,作为信号物质和微生物营养源调控着微生物的群落结构和多样性,而根区微生物区系的改变则反作用于植物的生长、发育和抗性。本文聚焦植物根系代谢物介导的植物-微生物互作,梳理了植物-微生物互作研究中次级代谢物的种类、作用及其检测手段;探讨了植物通过调节自身代谢物以适应品种进化及繁衍后代过程中发挥的功能作用;阐述了逆境胁迫下植物利用根系代谢物招募特异微生物(解磷、溶磷)或者有益微生物促进自身生长以缓解胁迫压力的机制;分析了根系代谢物作为信号物质诱导植物抗病的方式"求救假说",为可持续农业发展提供思路和理论依据。     

温带阔叶林中氮的保留机制：

 在过去的半个世纪中,森林生态系统物质能量流动模型的发展使得生态学者对矿物元素的来源、去处、转换速率及其干扰破坏的影响有了进一步的认识。其中“春坝”假设提出：在早春来临之际,林冠树木仍处于冬眠之时,林下地被植物先于林冠树木发芽开花,吸收土壤养分于植物体内,使得养分免于因雨水或雪水而造成的淋失而得以保留于森林生态系统中。本文对“春坝”假设的研究作一小结。现有的研究结果显示,早春地被植物对土壤氮的吸收量因不同生态系统而异;而土壤微生物对氮的吸收量远高于过去人们所认识的水平。早春地被物与土壤微生物都能在季节的变化中吸收土壤养分、共同起着“春坝”作用。两者所起作用大小因气候及生态系统的不同而有所改变。      

温带阔叶林中氮的保留机制:"春坝"假设及研究实例

在过去的半个世纪中,森林生态系统物质能量流动模型的发展使得生态学者对矿物元素的来源、去处、转换速率及其干扰破坏的影响有了进一步的认识.其中"春坝"假设提出在早春来临之际,林冠树木仍处于冬眠之时,林下地被植物先于林冠树木发芽开花,吸收土壤养分于植物体内,使得养分免于因雨水或雪水而造成的淋失而得以保留于森林生态系统中.本文对"春坝"假设的研究作一小结.现有的研究结果显示,早春地被植物对土壤氮的吸收量因不同生态系统而异;而土壤微生物对氮的吸收量远高于过去人们所认识的水平.早春地被物与土壤微生物都能在季节的变化中吸收土壤养分、共同起着"春坝"作用.两者所起作用大小因气候及生态系统的不同而有所改变.     

Effects of labile soil carbon on nutrient partitioning between an arctic graminoid and microbes

We measured partitioning of N and P uptake between soil microorganisms and potted Festuca vivipara in soil from a subarctic heath in response to factorial addition of three levels of labile carbon (glucose) combined with two levels of inorganic N and P. The glucose was added to either non-sterilized or sterilized (autoclaved) soils in quantities which were within the range of reported, naturally occurring amounts of C released periodically from the plant canopy. The aims were, firstly, to examine whether the glucose stimulated microbial nutrient uptake to the extent of reducing plant nutrient uptake. This is expected in nutrient-deficient soils if microbes and plants compete for the same nutrients. Secondly, we wanted to test our earlier␣interpretation that growth reduction observed in graminoids after addition of leaf extracts could be caused directly by labile carbon addition, rather than by phytotoxins in the extracts. Addition of high amounts of N did not affect the microbial N pool, whereas high amounts of added P significantly increased the microbial P pool, indicating a luxury P uptake in the microbes. Both plant N and in particular P uptake increased strongly in response to soil sterilization and to addition of extra N or P. The increased␣uptake led to enhanced plant growth when both elements were applied in high amounts, but only led to increased tissue concentrations without growth responses when the nutrients were added separately. Glucose had strong and contrasting effects on plant and microbial N and P uptake. Microbial N and P uptake increased, soil inorganic N and P concentrations were reduced and plant N and P uptake declined when glucose was added. The responses were dose-dependent within the range of 0–450 μg C g⁻¹ soil added to the non-sterilized soil. The opposite responses of plants and microbes showed that plant acquisition of limiting nutrients is dependent on release of nutrients from the soil microbes, which is under strong regulation by the availability and microbial uptake of labile C. Hence, we conclude, firstly, that the microbial populations can compete efficiently with plants for nutrients to an extent of affecting plant growth when the microbial access to labile carbon is high in nutrient deficient soils. We also conclude that reduced growth of plants after addition of leaf extracts to soil can be caused by carbon-induced shifts in nutrient partitioning between plants and microbes, and not necessarily by phytotoxins added with the extracts as suggested by some experiments. Received: 15 February 1997 / Accepted: 12 July 1997     

Endophyte roles in nutrient acquisition,root system architecture development and oxidative stress tolerance

Plants associate with communities of microbes (bacteria and fungi) that play critical roles in plant development, nutrient acquisition and oxidative stress tolerance. The major share of plant microbiota is endophytes which inhabit plant tissues and help them in various capacities. In this article, we have reviewed what is presently known with regard to how endophytic microbes interact with plants to modulate root development, branching, root hair formation and their implications in overall plant development. Endophytic microbes link the interactions of plants, rhizospheric microbes and soil to promote nutrient solubilization and further vectoring these nutrients to the plant roots making the soil-plant-microbe continuum. Further, plant roots internalize microbes and oxidatively extract nutrients from microbes in the rhizophagy cycle. The oxidative interactions between endophytes and plants result in the acquisition of nutrients by plants and are also instrumental in oxidative stress tolerance of plants. It is evident that plants actively cultivate microbes internally, on surfaces and in soils to acquire nutrients, modulate development and improve health. Understanding this continuum could be of greater significance in connecting endophytes with the hidden half of the plant that can also be harnessed in applied terms to enhance nutrient acquisition through the development of favourable root system architecture for sustainable production under stress conditions.     

Mineralization and distribution of nutrients in plants and microbes in four arctic ecosystems: responses to warming

Mineralization and nutrient distribution in plants and microbes were studied in four arctic ecosystems at Abisko, Northern Sweden and Toolik Lake, Alaska, which have been subjected to long-term warming with plastic greenhouses. Net mineralization and microbial immobilization were studied by the buried bag method and ecosystem pool sizes of C, N and P were determined by harvest methods. The highest amounts of organic N and P were bound in the soil organic matter. Microbial N and P constituted the largest labile pools often equal to (N) or exceeding (P) the amounts stored in the vegetation. Despite large pools of N and P in the soil, net mineralization of N and P was generally low during the growing season, except in the wet sedge tundra, and in most cases lower than the plant uptake requirement. In contrast, the microorganisms immobilized high amounts of nutrients in the buried bags during incubation. The same high immobilization was not observed in the surrounding soil, where the microbial nutrient content in most cases remained constant or decreased over the growing season. This suggests that the low mineralization measured in many arctic ecosystems over the growing season is due to increased immobilization by soil microbes when competition from plant roots is prevented. Furthermore, it suggests that plants compete well with microbes for nutrients in these four ecosystems. Warming increased net mineralization in several cases, which led to increased assimilation of nutrients by plants but not by the microbes.     

不同有机肥对烤烟根际土壤微生物的影响

通过田间小区试验,研究了不同有机肥(精制有机肥、生物有机肥)与减量20%化肥配施对烤烟根际微生物、青枯病防效及产量和品质的影响.结果表明: 与常规化肥(CF)相比,化肥减量20%配施有机肥(OF)或生物有机肥(BIO)均显著提高了烤烟根际的细菌数量和微生物总量,配施生物有机肥还显著提高了烤烟根际放线菌的数量,比OF增加44.3%,且真菌数量呈下降趋势.与CF处理相比,OF或BIO处理均显著提高了烤烟根际微生物利用碳底物的能力,BIO还显著提高了根际微生物利用酚类碳源的能力.OF和BIO处理均显著降低了烤烟青枯病的发生及危害程度,与CF相比,OF处理的烤烟青枯病发病率和病情指数分别下降了4%和8%,BIO处理的烤烟青枯病发病率和病情指数分别下降了23%和15.9%.OF和BIO处理均显著提高了烤烟的上等烟比例,比CF分别增加了10.5%和9.7%.BIO处理的产量和产值比OF分别提高17.1%和18.9%.     

植物-土壤反馈与草地群落演替:菌根真菌和土壤病原菌的调控作用

由植物引起的根际土壤生物或非生物环境的改变能够反馈影响群落中不同植物的生长,直接改变共存植物的相对竞争关系,推动群落结构的动态变化。作为土壤生物群落的重要组成部分,土壤微生物在植物-土壤反馈关系中起到重要的调控作用,对解释植物群落的演替进程和方向有着重要的意义。在草地植物群落演替的早期阶段和外来物种入侵的过程中,宿主植物对丛枝菌根真菌（AMF）的依赖性较低,受本地病原菌的影响较小,一般不存在负反馈。在演替后期,植物对AMF更具依赖性,而积累的病原菌则产生较强的负反馈效应,从而促进群落物种共存和植物多样性,提高草地生产力和稳定性。研究微生物-植物反馈机制不仅有助于完善草地退化与恢复理论,还对退化草地恢复治理的实践有着指导意义。未来关于根际微生物-植物反馈在草地群落演替中的作用应该加强以下几方面的研究：（1）在实验方法上,开展专性微生物-植物反馈研究;（2）在测定指标上,进一步量化不同微生物在反馈关系中的功能差异;（3）在研究对象上,加强土壤微生物在植物群落水平的反馈研究;（4）在应用上,明晰植物-土壤反馈在退化草地恢复过程中的作用,指导草地管理实践。     

根际微生物组组装与植物健康

土壤微生物拥有高度多样化的群落结构,其通过与植物发生复杂的相互作用影响植物健康,也被称为植物的第二基因组。最近研究表明植物能通过改变根际分泌物的组成影响根际微生物群落的组装,反之,根际微生物群落组成的改变能够通过影响植物营养吸收和抵御生物及非生物胁迫的能力影响植物健康。除此之外,农艺管理也是影响土壤微生物群落组装方式的重要因素。但到目前为止,根际微生物与宿主植物及土壤微生物之间互作机制的研究尚不清楚。本文将从农艺管理和宿主植物对微生物群落组装的影响及根际微生物组对植物健康的影响进行总结,为增加作物产量提供机会。     

Effect of long-term organic and mineral fertilization strategies on rhizosphere microbiota assemblage and performance of lettuce

Long-term agricultural fertilization strategies gradually change soil properties including the associated microbial communities. Cultivated crops recruit beneficial microbes from the surrounding soil environment via root exudates. In this study, we aimed to investigate the effects of long-term fertilization strategies across field sites on the rhizosphere prokaryotic (Bacteria and Archaea) community composition and plant performance. We conducted growth chamber experiments with lettuce (Lactuca sativa L.) cultivated in soils from two long-term field experiments, each of which compared organic versus mineral fertilization strategies. 16S rRNA gene amplicon sequencing revealed the assemblage of a rhizosphere core microbiota shared in all lettuce plants across soils, going beyond differences in community composition depending on field site and fertilization strategies. The enhanced expression of several plant genes with roles in oxidative and biotic stress signalling pathways in lettuce grown in soils with organic indicates an induced physiological status in plants. Lettuce plants grown in soils with different fertilization histories were visibly free of stress symptoms and achieved comparable biomass. This suggests a positive aboveground plant response to belowground plant–microbe interactions in the rhizosphere. Besides effects of fertilization strategy and field site, our results demonstrate the crucial role of the plant in driving rhizosphere microbiota assemblage.