丛枝菌根共生体的氮代谢运输及其生态作用

丛枝菌根真菌能与80%的陆生维管植物形成互惠共生关系,共生体的存在对促进植物营养吸收和提高抗逆性具有重要意义.丛枝菌根真菌从宿主植物获取其光合产物碳水化合物的同时,通过外生菌丝吸收各种氮源,有效增强了宿主植物对氮素的吸收,以及氮在植物居群和群落水平上的交流,改善了植物营养代谢,增强了植物应对外界环境胁迫的能力.而共生体对氮的吸收、转运,以及氮从真菌到宿主植物的传输、代谢机制至今仍有许多问题亟待解决.本文综述了当前丛枝菌根共生体中氮传输代谢的主要机制,以及碳、磷对共生体氮传输代谢的影响;从群落和生态系统水平,简要阐述了丛枝菌根真菌在植物中氮分配的作用和对宿主植物的生态学意义,并提出共生体中氮代谢的一些需要深入研究的问题.
     

植物内生菌群落组成及其功能研究进展

植物内生菌是栖居在植物组织内部、以宿主植物代谢物为营养物质的一类微生物。植物内生菌种类丰富,并通过不同的方式定殖在植物体内,与宿主植物互利共生。植物内生菌促进植物的生长发育,通过增强营养物质吸收、与病原菌竞争生态位、在代谢过程中产生抗菌物质以及诱导宿主植物产生抗性等机制,提高宿主植物的抗逆性,并且其群落的组成因生态环境、寄主植物不同部位、生长周期的不同发育阶段而显著不同。人们利用传统组织分离法、高通量测序法以及人工重组生物群落等研究方法对植物与内生菌相互作用进行了深入的研究。本文对植物内生菌群落的组成、内生菌对宿主植物影响的功能研究以及人工内生菌群的构建进行重点阐述,以期为植物内生菌的开发和利用奠定坚实基础。     

药用植物内生菌对宿主植物促生作用机制研究进展

植物内生菌是一类能够代谢产生新颖生物分子和多种酶类的重要微生物资源,在农业、植保、制药等领域具有广阔的应用前景。了解内生菌与药用植物间相互关系是当前促进药用植物生长和提升药材品质的重要途径。植物内生菌资源具有丰富的多样性,对宿主植物生长发挥着重要功能,如固氮、溶磷、产生铁载体、分泌植物激素吲哚乙酸(indole-3-acetic acid,IAA)、产生ACC脱氨酶(1-aminocyclopropane-1-carboxylate deaminase)、增强宿主抗逆性、产生次生代谢产物等。本文通过相关文献回顾,聚焦内生菌与药用植物间的关系,着重探讨药用植物内生菌对宿主植物的促生作用机制,展望新技术在植物内生菌研究方面的应用,以便有效利用分子手段阐明内生菌对药用植物的促生长作用,为其在相关领域的应用提供理论参考。     

丛枝菌根共生体中碳、氮代谢及其相互关系

丛枝菌根共生体(arbuscular mycorrhiza, AM)是丛枝菌根真菌(arbuscular mycorrhizal fungi, AMF)与宿主植物之间形成的互惠共生形式.共生体中的碳、氮交换和代谢影响着宿主植物和共生真菌之间的营养平衡和资源重新分配,在物质和能量循环中发挥着重要作用.宿主植物光合固定的碳输送到真菌内,并且分解和释放真菌所需的生命物质和能量,包括促进孢子萌发、菌丝生长和提高氮等营养元素的吸收;而菌根真菌利用宿主植物提供的碳骨架和能量,发生氮的转化和运输,最终传递给宿主植物供其利用.本文综述了丛枝菌根共生体中碳、氮传输和代谢的主要模式,碳、氮的交互影响和调控机制,以促进丛枝菌根在可持续农业和生态系统中的应用.     

外来入侵植物的氮代谢及其土壤氮特征

研究了4种外来入侵植物(五爪金龙、南美蟛蜞菊、金腰箭和马缨丹)和1种本地植物鸡矢藤(对照)的氮代谢及其土壤氮特征.结果表明:外来人侵植物的组织硝酸还原酶活性、根际土壤NH4-N、NO3-N含量、蛋白酶活性和脲酶活性均较高,分别为鸡矢藤的1.65～4.34、1.56～2.15、1.72～3.11、1.43～3.23和1.41～3.33倍,而植物组织硝态氮含量则较低,仅为鸡矢藤的17.5%～50.6%.相关分析表明:植物组织硝酸还原酶活性与根际土壤总氮、NH4-N、NO3-N含量呈显著正相关(P<0.05),与蛋白酶活性和脲酶活性呈极显著正相关(P<0.01).这说明,外来植物入侵使土壤氮代谢加快,氮的生物有效性增强,氮同化能力提高,并且较好地将植物体氮素代谢与土壤氮素代谢协调起来.因此,较强的氮素同化能力与加速土壤氮素的转化可能是植物成功入侵的重要机制之一.     

氮代谢参与植物逆境抵抗的作用机理研究进展

近年来,植物所受到的诸如干旱、盐、高温、低氧、重金属胁迫和营养元素缺乏等环境胁迫越来越多,严重影响了植物的生长发育及作物的质量和产量。氮素是植物生长发育所需的必需营养元素,同时也是核酸、蛋白质和叶绿素的重要组成成分,其代谢过程与植物抵抗逆境的能力息息相关。氮代谢是指植物对氮素的吸收、同化和利用的全过程,是植物体内基础代谢途径之一。氮代谢主要从氮素吸收、同化及氨基酸代谢等方面参与植物的抗逆性,并通过调节离子吸收和转运、稳定细胞形态和蛋白质结构、维持激素平衡和细胞代谢水平、减少体内活性氧(reactive oxygen species,ROS)生成以及促进叶绿素合成等生理机制来影响植物抵抗非生物胁迫的能力。因此,提高植物在逆境下的氮代谢水平是减轻外界胁迫对其损伤的一种潜在途径。该文从氮素同化的基本途径出发,分别阐述了氮代谢在干旱胁迫、盐胁迫和高温胁迫等多个方面的逆境抵抗过程中的作用机理,为氮代谢参与植物抗逆性研究提供了有利参考。     

不同氮水平下内生固氮菌对入侵植物南美蟛蜞菊生长策略的影响

很多因素可以影响外来植物的入侵性,其中内生菌可能扮演了极其重要的角色。为了探究内生固氮菌对入侵植物生长策略的影响,本研究分别在高和低两种氮水平下,种植接种和不接种内生固氮菌的入侵植物南美蟛蜞菊(Sphagneticola trilobata)和同属本地植物蟛蜞菊(S. calendulacea),对比研究它们的生长和氮含量。从南美蟛蜞菊茎段分离出内生固氮菌Kosakonia sp. WTB-JS007,发现其对南美蟛蜞菊和蟛蜞菊这两种植物生长策略的影响存在显著差异,且这种差异不受氮水平的影响。在低氮处理下,接种WTB-JS007对蟛蜞菊的生长及植株氮含量没有显著影响,但显著提高了南美蟛蜞菊的地上生物量(30.48%),促进了匍匐茎的伸长,减少了地下生物量(56.58%),并提高了植株总氮含量(47.51%)。类似地,在高氮水平下,内生固氮菌亦显著影响南美蟛蜞菊地上部分的生长,但对蟛蜞菊生长的影响不显著。这些结果表明,内生固氮菌对入侵和非入侵植物生长、生物量分配及氮获取的影响存在显著差异,而这种生长策略的差异有利于入侵植物地上部分的快速生长与扩张,从而提高其入侵性。     

丛枝菌根真菌在土壤氮素循环中的作用

作为植物需求量最大的营养元素,氮素是陆地生态系统初级生产力的主要限制因子。丛枝菌根真菌能与地球上80%以上的陆生植物形成菌根共生体,帮助宿主植物吸收土壤中的P、N等矿质养分。目前,丛枝菌根真菌与氮素循环相关研究侧重于真菌对氮素的吸收形态以及共生体中氮的传输代谢机制,却忽略了丛枝菌根真菌在固氮过程、矿化与吸收过程、硝化过程、反硝化过程以及氮素淋洗过程等土壤氮素循环过程中所起到的潜在作用,并且越来越多的证据也表明丛枝菌根真菌是影响土壤氮素循环过程的重要因子。总结了丛枝菌根真菌可利用的氮素形态及真菌的氮代谢转运相关基因的研究现状;重点分析了丛枝菌根真菌在调控土壤氮素循环过程中的潜在作用以及在生态系统中的重要生态学意义,同时提出了丛枝菌根真菌在土壤氮素循环过程中一些需要深入研究的问题。     

植物内生菌是有待深入开发的资源宝库

植物内生菌是指那些在其生活史的一定阶段或者全部阶段生活于健康植物各种组织和器官内部的真菌或者细菌,被感染的宿主植物(至少是暂时)不表现出外在症状。在长期的协同进化过程中,大部分植物内生菌都与植物形成了互惠互利的关系,在从宿主那里获得稳定的生活环境的同时,可增强或赋予宿主抗病、抗干旱、固氮等能力,或通过其代谢产物促进植物的生长;有些内生菌还被发现能够产生与宿主相同或相似的活性物质。因此对植物内生菌的研究从20     

外源输入氮的有效性及形态对植物生长与生理影响的研究进展

氮(N)是控制植物结构和功能以及维持生态系统稳定的重要营养元素之一,外源输入氮素的有效性及形态的差异对植物的生长发育和生理特征产生显著的影响。由于全球气候变化和人类活动的干扰,大气氮沉降量日益增加,氮形态也发生改变,严重破坏植物的正常生长和生态系统的平衡稳定,已成为研究学者关注的热点问题。本文综述了不同氮输入水平和形态对植物的生长、光合作用、养分吸收以及代谢酶活性等方面的影响,归纳出：(1)适量氮输入能促进植物生长、光合作用和养分吸收能力,但当超出植物承受的阈值后,则对其产生抑制作用;(2)由于植物对氮素形态吸收偏好的差异,铵态氮(NH₄⁺-N)和硝态氮(NO₃^--N)对植物的生长、光合作用、养分吸收以及代谢酶活性的影响效果不同,且适宜铵硝配比相较于单一某种氮素添加对大多数植物的促进作用更显著。提出未来的研究方向应着重考虑4个方面：(1)开展大尺度的长期监测控制实验;(2)利用分子生物学技术深入探究氮形态对植物影响的微观机理;(3)重点关注土壤根际环境对植物根系氮素吸收的影响;(4)综合分析氮输入与...     

A consortium of non‐rhizobial endophytic microbes from Typha angustifolia functions as probiotic in rice and improves nitrogen metabolism

• Endophytic microbes isolated from plants growing in nutrient‐deficient environments often possess properties that improve nutrition of agriculturally important plants. A consortium of non‐rhizobial endophytic microbes isolated from a macrophyte Typha angustifolia growing in the marginal wetlands associated with a Uranium mine was characterized for their beneficial effect on rice and the mechanisms of growth promotion were investigated.
• The microbes were identified and characterized for their potential plant growth promoting (PGP) properties. Effect of these microbes on nitrogen (N)‐metabolism of rice was tested as Typha endophytes were predominantly (N)‐fixing. Relative N‐use efficiency and expression of genes involved in N‐uptake and assimilation were investigated in treated plants.
• Evidence of horizontal gene transfer (HGT) of dinitrogen reductase gene was observed within the consortium from a Pseudomonas stutzeri strain. The consortium behaved as plant probiotic and showed substantial growth benefits to Typha, their natural host as well as to rice. Typha endophytes colonized rice endosphere significantly increasing biomass, shoot length and chlorophyll content in rice plants both under N‐sufficient and N‐deficient conditions. N‐uptake and assimilation genes were upregulated in plants treated with the endophytes even after three weeks post infection.
• Our results suggested, HGT of nitrogen‐fixation trait to be highly prevalent among endophytes isolated from nutrient‐poor habitats of the uranium mine. A long‐term nitrogen deficiency response in the treated plants was elicited by the consortium improving N‐uptake, assimilation and relative N‐use efficiency of rice plants. This appeared to be at least one of the main strategies of plant growth promotion.

     

Pea aphid promotes amino acid metabolism both in Medicago truncatula and bacteriocytes to favor aphid population growth under elevated CO2

Rising atmospheric CO₂ levels can dilute the nitrogen (N) resource in plant tissue, which is disadvantageous to many herbivorous insects. Aphids appear to be an exception that warrants further study. The effects of elevated CO₂ (750 ppm vs. 390 ppm) were evaluated on N assimilation and transamination by two Medicago truncatula genotypes, a N‐fixing‐deficient mutant (dnf1) and its wild‐type control (Jemalong), with and without pea aphid (Acyrthosiphon pisum) infestation. Elevated CO₂ increased population abundance and feeding efficiency of aphids fed on Jemalong, but reduced those on dnf1. Without aphid infestation, elevated CO₂ increased photosynthetic rate, chlorophyll content, nodule number, biomass, and pod number for Jemalong, but only increased pod number and chlorophyll content for dnf1. Furthermore, aphid infested Jemalong plants had enhanced activities of N assimilation‐related enzymes (glutamine synthetase, Glutamate synthase) and transamination‐related enzymes (glutamate oxalate transaminase, glutamine phenylpyruvate transaminase), which presumably increased amino acid concentration in leaves and phloem sap under elevated CO₂. In contrast, aphid infested dnf1 plants had decreased activities of N assimilation‐related enzymes and transmination‐related enzymes and amino acid concentrations under elevated CO₂. Furthermore, elevated CO₂ up‐regulated expression of genes relevant to amino acid metabolism in bacteriocytes of aphids associated with Jemalong, but down‐regulated those associated with dnf1. Our results suggest that pea aphids actively elicit host responses that promote amino acid metabolism in both the host plant and in its bacteriocytes to favor the population growth of the aphid under elevated CO₂.     

Properties of bacterial endophytes and their proposed role in plant growth

Bacterial endophytes live inside plants for at least part of their life cycle. Studies of the interaction of endophytes with their host plants and their function within their hosts are important to address the ecological relevance of endophytes. The modulation of ethylene levels in plants by bacterially produced 1-aminocyclopropane-1-carboxylate deaminase is a key trait that enables interference with the physiology of the host plant. Endophytes with this capacity might profit from association with the plant, because colonization is enhanced. In turn, host plants benefit by stress reduction and increased root growth. This mechanism leads to the concept of 'competent' endophytes, defined as endophytes that are equipped with genes important for maintenance of plant-endophyte associations. The ecological role of these endophytes and their relevance for plant growth are discussed here.     

Endophyte-assisted phytoremediation: mechanisms and current application strategies for soil mixed pollutants

Abstract

Phytoremediation uses plants and associated microbes to remove pollutants from the environment and is considered a promising bioremediation method. Compared with well-described single contaminant treatments, the number of studies reporting phytoremediation of soil mixed pollutants has increased recently. Endophytes, including bacteria and fungi, exhibit beneficial traits for the promotion of plant growth, stress alleviation, and biodegradation. Moreover, endophytes either directly or indirectly assist host plants to survive high concentrations of organic and inorganic pollutants in the soil. Endophytic microorganisms can also regulate the plant metabolism in different ways, exhibiting a variety of physiological characteristics. This review summarizes the taxa and physiological properties of endophytic microorganisms that may participate in the detoxification of contaminant mixtures. Furthermore, potential biomolecules that may enhance endophyte mediated phytoremediation are discussed. The practical applications of pollutant-degrading endophytes and current strategies for applying this valuable bio-resource to soil phytoremediation are summarized.     

Fungal endophytes: common host plant symbionts but uncommon mutualists

Fungal endophytes are extremely common and highly diverse microorganismsthat live within plant tissues, but usually remain asymptomatic.Endophytes traditionally have been considered plant mutualists,mainly by reducing herbivory via production of mycotoxins, suchas alkaloids. However, the vast majority of endophytes, especiallyhorizontally-transmitted ones commonly found in woody plants,apparently have little or no effect on herbivores. For the systemic,vertically-transmitted endophytes of grasses, mutualistic interactionsvia increased resistance to herbivores and pathogens are morecommon, as predicted by evolutionary theory. However, even inthese obligate symbioses, endophytes are often neutral or evenpathogenic to the host grass, depending on endophyte and plantgenotype and environmental conditions. We present a graphical model based upon variation in nitrogenflux in the host plant. Nitrogen is a common currency in endophyte/hostand plant/herbivore interactions in terms of limitations tohost plant growth, enhanced uptake by endophytes, demand forsynthesis of nitrogen-rich alkaloids, and herbivore preferenceand performance. Our graphical model predicts that low alkaloid-producingendophytes should persist in populations when soil nutrientsand herbivory are low. Alternatively, high alkaloid endophytesare favored under increasing herbivory and increasing soil nitrogen,at least to some point. At very high soil nitrogen levels, uninfectedplants may be favored over either type of infected plants. Thesepredictions are supported by patterns of infection and alkaloidproduction in nature, as well by a manipulative field experiment.However, plant genotype and other environmental factors, suchas available water, interact with the presence of the endophyteto influence host plant performance.     

植物内生菌及其防治植物病害的研究进展

综述了植物内生菌及其防治植物病害的研究进展.植物内生菌分布广,种类多,几乎存在于所有目前已研究过的陆生及水生植物中,目前全世界至少已在80个属290多种禾本科植物中发现有内生真菌,在各种农作物及经济作物中发现的内生细菌已超过120种.感染内生菌的植物宿主往往具有生长快速、抗逆境、抗病害、抗动物危害等优势,比未感染内生菌的植株更具生存竞争力.植物内生菌的防病机理主要表现在通过产生抗生素类,水解酶类,植物生长调节剂和生物碱类物质,与病原菌竞争营养物质,增强宿主植物的抵抗力以及诱导植物产生系统抗性等途径抑制病原菌生长.另外,对植物内生真菌和内生细菌的分离、筛选和检测方法;利用植物内生菌控制植物病害的途径如人工接种内生菌,利用内生菌代谢产生的抗生素以及将内生菌作为基因工程的载体菌等进行了综述.同时,对植物内生菌作为生物防治因子未来发展前景及存在的问题进行了讨论.利用植物内生菌作为生物防治因子进行大田防病,需要考虑它的病理学、生态学和形态学等方面的影响.     

Carbon and nitrogen metabolism in legume root nodules

The literature concerning the metabolism of carbon compounds during the reduction, assimilation and translocation of nitrogen in root nodules of leguminous plants is reviewed. The reduction of dinitrogen requires an energy source (ATP) and a reluctant which are both supplied by respiratory catabolism of carbohydrates produced by the host plant. Photosynthates are also required to generate the carbon skeletons for amino acid or urcide synthesis during the assimilation of ammonia produced by the bacteria within the nodule tissue. Competition for photosynthates occurs between the bacteroids, nodule tissue and the various vegetative and reproductive sinks in the host plant. The nature of carbon compounds involved in these processes, their routes of metabolism, the mechanisms of control and the partitioning of metabolises between the various sites of utilization are only poorly understood. It is apparent that dinitrogen is reduced to ammonia in the bacteroids. Both fast- and slow-growing strains of Rhizobium possess the Entner-Doudoroff pathway of glucose catabolism, and some, if not all, enzymes of the Emden-Meyerhof pathway. Some bacterial cultures also metabolize carbon through the ketogluconate pathway but only the fast-growing strains of cultured rhizobia possess the key enzyme of the pentose phosphate pathway (6-phosphogluconate dehydrogenase). The host cells are thought to contain the complete Emden-Meyerhof pathway and tricarboxylic acid cycle, which provides the carbon skeletons for assimilation of the ammonia, formed by the bacteroids, into α-amino acids. A pathway of anapleurotic carbon conservation, operative in the host cells, synthesizes oxaloacetic acid through β-carboxylation of phosphoenol pyruvate. This process could be important in the recapture and assimilation of respired CO₂ in the rhizosphere. The main route of assimilation of ammonia produced by the bacteroids would appear to be via the glutamine synthetase-glutamate synthase pathway in the host cells. However, glutamate dehydrogenase may also be involved in ammonia assimilation. These enzymes also occur in in vitro cultures of Rhizobium and in bacteroids where they presumably participate in the synthesis of amino acids for growth of the bacteria or bacteroids. Nitrogen assimilated into glutamine or glutamate is exported from the nodules in a variety of forms, which include asparagine, glutamine, aspartate, homoserine and allantoates, in proportions which depend on the legume species. Studies on regulation of the overall process have focussed on expression of bacteroid genes and on the control of enzyme activity, at the level of nitrogenase and enzymes of nitrogen assimilation in particular. However, due to the wide range of experimental techniques, environmental conditions and plant species which have been used, no clear conclusions can yet be drawn. The pathways of carbon flow in nitrogen metabolism, particularly in relation to the synthesis of ureides and the regulation of carbon metabolism, remain key areas for future research in symbiotic nitrogen fixation.     

水分胁迫下内生真菌感染对黑麦草叶内游离脯氨酸和脱落酸含量的影响

以黑麦草为实验对象,研究了干旱胁迫条件下内生真菌感染对植株叶片含水量和叶内游离脯氨酸含量的影响,同时对渗透胁迫条件下植株叶内ABA含量的变化进行了分析。结果表明：①内生真菌的感染有助于使叶片保持较高的含水量;②在两种形式的水分胁迫下,。前期至中期高感染种群的叶片游离脯氨酸含量低于感染种群,而在末期则有高出低感染种群的趋势;③内生真菌感染对黑麦草叶内ABA累积的正效应只发生在轻度渗透胁迫下的较短时间范围内。     

Ureide metabolism in higher plants

The synthesis, transport and assimilation of the ureides, allantoin and allantoic acid, in higher plants is reviewed. Evidence indicates that in nodulated legumes ureides are synthesized from products of N₂-fixation via purine synthesis and degradation. Their synthesis in other plants also appears to be via purine degradation but is dependent on the inorganic nitrogen source fed to the plant; greatest ureide production is associated with ammonium assimilation. The use of ureides rather than amides for N-transport from the root to the shoot via the xylem stream results in an improved carbon economy of the plant. Good evidence for the transport of ureides in the phloem is lacking for most species examined although it is assumed to be important, particularly in fruit and seed development. Ureides are stored and assimilated mainly in the shoot. The precise pathways, localization and regulation of ureide assimilation are poorly understood and require further investigation. Similarities exist between the properties of the enzymes involved in ureide assimilation in higher plants and in micro-organisms. However, the evidence that light appears to be involved in ureide assimilation in green tissues suggests that different regulatory mechanisms may exist in plants compared with micro-organisms. The economically important legume crops such as soybeans, cowpeas and Phaseolus sp. are all ureide producers. To aid our understanding of the productivity of these plants knowledge of how ureide-N is converted into seed protein is essential.     

Leaf nitrogen dioxide uptake coupling apoplastic chemistry, carbon/sulfur assimilation, and plant nitrogen status

Emission and plant uptake of atmospheric nitrogen oxides (NO + NO₂) significantly influence regional climate change by regulating the oxidative chemistry of the lower atmosphere, species composition and the recycling of carbon and nutrients, etc. Plant uptake of nitrogen dioxide (NO₂) is concentration-dependent and species-specific, and covaries with environmental factors. An important factor determining NO₂ influx into leaves is the replenishment of the substomatal cavity. The apoplastic chemistry of the substomatal cavity plays crucial roles in NO₂ deposition rates and the tolerance to NO₂, involving the reactions between NO₂ and apoplastic antioxidants, NO₂-responsive germin-like proteins, apoplastic acidification, and nitrite-dependent NO synthesis, etc. Moreover, leaf apoplast is a favorable site for the colonization by microbes, which disturbs nitrogen metabolism of host plants. For most plant species, NO₂ assimilation in a leaf primarily depends on the nitrate (NO₃ ⁻) assimilation pathway. NO₂–N assimilation is coupled with carbon and sulfur (sulfate and SO₂) assimilation as indicated by the mutual needs for metabolic intermediates (or metabolites) and the NO₂-caused changes of key metabolic enzymes such as phosphoenolpyruvate carboxylase (PEPc) and adenosine 5′-phosphosulfate sulfotransferase, organic acids, and photorespiration. Moreover, arbuscular mycorrhizal (AM) colonization improves the tolerance of host plants to NO₂ by enhancing the efficiency of nutrient absorption and translocation and influencing foliar chemistry. Further progress is proposed to gain a better understanding of the coordination between NO₂–N, S and C assimilation, especially the investigation of metabolic checkpoints, and the effects of photorespiratory nitrogen cycle, diverse PEPc and the metabolites such as cysteine, O-acetylserine (OAS) and glutathione.