丛枝菌根真菌-植物共生体耐盐机制的研究进展

土壤盐碱化严重制约农业发展并影响生态环境.丛枝菌根真菌(AMF)与植物形成的共生体作为生态系统的有机组成部分,因其形成的广泛性,可增强植物抗盐碱胁迫的能力,具有不可忽视的生态调节作用.本文从盐胁迫对AMF发育的影响、盐胁迫下AMF对植物生长的影响、AMF增强植物耐盐性的内在机制等3个方面阐述了丛枝菌根真菌-植物共生体耐盐性的机制.并结合当前研究存在的难题以及发展趋势对今后本领域的研究方向做出展望.     

丛枝菌根共生的信号转导及其相关基因

大多数植物根系能够与某些真菌形成相互依存、互惠互利的菌根共生关系.植物在提供给丛枝菌根真菌赖以生存的碳源的同时,也通过真菌从土壤中吸取矿质营养.丛枝菌根能够促进植物生长,提高植物抗逆性和抵御外界不良环境,对提高农林业生产效率、增强生态系统稳定性及维护生物多样性具有重要的意义.菌根的形成是一系列信号分子交换传递和共生相关基因表达调控的结果.在信号转导途径中,共生受体样蛋白激酶、离子通道和钙/钙调依赖性蛋白激酶基因的表达对菌根的形成是不可或缺的.就丛枝菌根共生的信号转导机制以及信号途径中3个必需基因的结构、功能及研究现状进行了综述.     

丛枝菌根（AM）生物技术在现代农业体系中的生态意义

菌根是植物根系与特定的土壤真菌形成的共生体,有利于生态系统中养分循环,协助植物抵御不良环境胁迫．自然条件下,大多数植物表现一定的菌根依赖性,在植株根系发育过程中如能与适宜的菌根真菌形成良好的菌根结构,可提高产量,改善品质,其中丛枝菌根是最普遍的类型．丛枝菌根帮助植物抵御不良环境胁迫及病虫害,促进植物健康生长,可减少化学肥料、杀虫剂施用量,以减少对环境、生态不利的化学物质施用量．丛枝菌根共生体可加速根系生长,提高对移动性低的无机离子吸收,加速养分循环利用,增强植物对不良胁迫(生物与非生物)因素的耐受力,形成良好的土壤结构,提高植物群体的多样性．文章综述了丛枝菌根真菌生态特征,丛枝菌根对寄主植物的影响,丛枝菌根生物技术应用于农业体系的生态意义及其应用潜力．     

丛枝菌根对植物根际逆境的生态学意义

近年来,我国在菌根分子生物学、菌根营养学、菌根分类学和菌根生态学等方面取得了令人瞩目的研究成果,其中对丛枝菌根真菌(AMF)的研究居多。AMF能与大部分陆地植物根系形成共生关系,促进植物生长发育,提高植物抗逆性,在保持生态平衡、保护生态环境等方面发挥重要作用。本文主要从非生物胁迫(干旱胁迫、重金属污染、盐碱胁迫)和生物胁迫(致病菌和线虫侵染)方面介绍了AMF在植物根际逆境中发挥的生态功能及作用机制,提出了该研究领域尚存的不足之处和研究前景,为AMF后续研究提供参考。     

丛枝菌根形成过程及其信号转导途径

丛枝菌根是由一类土壤中古老的丛枝菌根真菌与植物根系形成的互利互惠共生体。通过共生作用丛枝菌根真菌帮助宿主植物提高水和矿质营养（特别是磷）的吸收效率。作为回报,大约20%的光合作用产物被转移到丛枝菌根真菌中,供其完成自身的生活史。丛枝菌根形成的过程中,需要植物与丛枝菌根真菌之间进行一系列信号分子的识别、交换以及信号转导作用,这一过程由一系列植物和菌根真菌的基因控制。首先,植物会分泌一种植物激素——独角金内酯来诱导菌根真菌加速分支,而菌根真菌也会分泌脂质几丁寡糖促进植物与其形成菌根。加速分支的菌根真菌接触到植物根部以后,会附着在植物根的表皮并形成附着胞,通过附着胞穿透植物根的表皮,最后进入维管组织附近的皮层细胞并在其中不断进行二叉分支,形成特有的丛枝结构。通过对模式植物共生现象的研究,已经发现很多植物基因参与到共生形成的信号转导过程中,包括早期植物反应的基因、菌根与根瘤共生共同需要的转导因子以及菌根特异的信号分子等。本文对菌根的形成过程及信号转导途径进行详细的介绍,为人们深入研究菌根关系提供参考。     

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.
     

丛枝菌根共生关系的信号机制研究进展

在植物与微生物的共生体中,最广泛的互惠共生体就是丛枝菌根.真菌在植物根系形成菌根后,菌丝通过根的皮层细胞获取植物提供的碳源,同时将矿物营养和水从土壤转运到皮层细胞,这种共生过程的研究在生物多样性的保护、陆生植物的起源与演化、退化生态系统的修复与重建以及农业、林业和园艺业的应用具有重要的意义.近年来丛枝菌根真菌与植物根系建立共生关系的信号传导途径和作用机制备受关注,也取得了突破性的进展.本文对丛枝菌根真菌与植物根系在共生关系形成、营养交换以及防御方面的分子信号和细胞方面的研究进展进行综述,并对发展前景作以展望.     

Arbuscular mycorrhizal fungi protect a subtropical tree species exposed to simulated acid rain by accelerating photosynthetic ability,antioxidant enzymes and osmolyte accumulation

丛枝菌根真菌对其宿主光合能力、抗氧化酶和渗透物质积累的促进作用 及其抗酸雨机制的探讨 酸雨在中国南方发生频繁,对亚热带树种生长具有明显抑制作用。以往研究表明,丛枝菌根真菌(AM真菌)可以缓解酸雨对宿主植物的胁迫效应。榉树(Zelkova serrata)为中国南方主要经济树种之一,其如何与共生AM真菌协同、增强其抗酸雨胁迫的能力是本项研究所要探讨的关键科学问题。通过温室控制实验,将榉树幼苗随机接受4个水平的AM真菌接种处理(接种灭菌菌种;单独接种Rhizophagus intraradices;单独接种Diversispora versiformis;接种这两种菌种的混合菌种)和3个pH水平(pH2.5、pH4.0和pH5.6)的硫酸型酸雨和硝酸型酸雨处理组成的12个处理组合,同时测定其生长、光合性能、抗氧化酶、渗透调节和土壤酶的响应格局。研究发现酸雨处理显著降低了非菌根榉树幼苗的总干重、总叶绿素含量、叶片净光合速率和可溶性蛋白的含量;接种AM真菌,特别是接种混合菌种,显著提高了强酸胁迫下榉树幼苗的总干重、光合性能、丙二醛、过氧化物酶、超氧化物歧化酶、可溶性蛋白和根系酸性磷酸酶活性。此外,菌根效应依赖于AM真菌的种类和酸胁迫的梯度。本研究 结果表明,AM真菌对榉树幼苗抗酸胁迫的调控作用主要源于调节宿主植株光合能力、抗氧化酶和渗透物质的积累。榉树与其共生AM真菌在应对酸胁迫上协同机制的解析为该树种在中国南方酸雨区的栽培提供理论基础、具有重要的实践指导意义。     

丛枝菌根真菌与植物硫素营养研究进展

丛枝菌根真菌是土壤微生物群落的重要组成部分,是最常见的地下共生菌,对植物和土壤具有多种有益作用。本文阐述了近年来丛枝菌根真菌对植物吸收土壤硫素的最新进展,在目前耕地缺硫状况下,着重分析了丛枝菌根真菌改善植物硫素营养以及丛枝菌根真菌利用硫素的分子调控机制,总结了影响菌根硫代谢的因素,并指出该研究方向仍存在的一些问题以及未来的研究侧重点。     

丛枝菌根真菌与植物共生对植物水分关系的影响及机理

自1885年Frank首次提到菌根(mykorhiza)概念以来,大量的试验证实了丛枝菌根真菌(AMF)与植物根系之间形成具有一定结构和功能的共生体,促进植物生长并提高干旱耐受能力,在干旱生态系统中发挥重要的作用。该研究多集中在对宿主植物生理生态的影响及其机制方面,然而菌根共生对宿主植物水分吸收和信号产生、传递的影响研究少而分散,缺少系统总结。综述了最近四十多年丛枝菌根真菌与植物共生体对宿主植物干旱适应性影响研究进展,讨论了菌根共生对植物根冠通讯的影响及机理。干旱胁迫下AMF与植物共生,通过影响宿主植物一系列生理生态过程,提高宿主植物横向根压和纵向蒸腾拉力。经典的Ohm吸水模型是该方向最有代表性的研究成果,该模型揭示了菌根共生的根外菌丝具有不同于根细胞的细胞结构和水分运输性能,这为宿主植物提供一种特殊的快速吸水方式,可提高植物对土壤水分的吸收和运输能力。研究表明,AMF会影响宿主植物根冠通讯过程,如诱发信号级联反应,诱导根系尽早感知水分胁迫并产生非水力根源信号,提高宿主对干旱的耐受性。讨论了AMF在根冠通讯分子机制研究方面存在的问题及可能的解决途径,展望了AMF在干旱农业生产中的应用潜力。     

不同强度盐胁迫下AM真菌对羊草生长的影响

不同浓度NaCl盐处理下,AM真菌对羊草(Leymus chinensis)的侵染能力和对植物生长的影响,从植物形态和离子含量角度探讨了AM真菌提高羊草耐盐性的作用机理。结果表明,在高盐胁迫下,AM真菌显著降低了盐胁迫效应,提高了羊草生物量,菌根效应明显。菌根化羊草的根茎比显著增加,并且N、P浓度较高,Na~+和Cl~-离子浓度较低,表明AM真菌即促进羊草对营养元素的吸收,又减少了离子毒害。菌根化羊草的Ca~(2+)和K~+离子浓度,以及P/Na~+和K~+/Na~+比高于非菌根化羊草,表明AM真菌可通过调节渗透势以避免或减缓盐胁迫造成的生理缺水。随着盐胁迫的增加,菌根化羊草对磷的依赖性逐渐转换为对钾的依赖性。研究结果有助于揭示AM真菌提高植物耐盐能力的作用机理,并对应用菌根技术修复盐化草地具有理论指导意义。     

Antioxidative defense under salt stress

Salt tolerance is a complex trait involving the coordinated action of many gene families that perform a variety of functions such as control of water loss through stomata, ion sequestration, metabolic adjustment, osmotic adjustment and antioxidative defense. In spite of the large number of publications on the role of antioxidative defense under salt stress, the relative importance of this process to overall plant salt tolerance is still a matter of controversy. In this article, the generation and scavenging of reactive oxygen species (ROS) under normal and salt stress conditions in relation to the type of photosynthesis is discussed. The CO₂ concentrating mechanism in C4 and CAM plants is expected to contribute to decreasing ROS generation. However, the available data supports this hypothesis in CAM but not in C4 plants. We discuss the specific roles of enzymatic and non enzymatic antioxidants in relation to the oxidative load in the context of whole plant salt tolerance. The possible preventive antioxidative mechanisms are also discussed.Key words: salt stress, generation of ROS, type of photosynthesis, scavenging of ROS, preventive antioxidative defense     

Plant salt-tolerance mechanism: A review

Almost all crops that are important to humans are sensitive to high salt concentration in the soil. The presence of salt in soil is one of the most significant abiotic stresses in farming. Therefore, improving plant salt tolerance and increasing the yield and quality of crops in salty land is vital. Transgenic technology is a fast and effective method to obtain salt-tolerant varieties. At present, many scholars have studied salt damage to plant and plant salt-tolerance mechanism. These scholars have cloned a number of salt-related genes and achieved high salt tolerance for transgenic plants, thereby showing attractive prospects.In this paper, the salt-tolerance mechanism of plants is described from four aspects: plant osmotic stress, ion toxicity, oxidative stress, and salt tolerance genes. This review may help in studies to reveal the mechanism of plant salt tolerance, screen high efficiency and quality salt tolerance crops.     

Current developments in arbuscular mycorrhizal fungi research and its role in salinity stress alleviation: a biotechnological perspective

Arbuscular mycorrhizal fungi (AMF) form widespread symbiotic associations with 80% of known land plants. They play a major role in plant nutrition, growth, water absorption, nutrient cycling and protection from pathogens, and as a result, contribute to ecosystem processes. Salinity stress conditions undoubtedly limit plant productivity and, therefore, the role of AMF as a biological tool for improving plant salt stress tolerance, is gaining economic importance worldwide. However, this approach requires a better understanding of how plants and AMF intimately interact with each other in saline environments and how this interaction leads to physiological changes in plants. This knowledge is important to develop sustainable strategies for successful utilization of AMF to improve plant health under a variety of stress conditions. Recent advances in the field of molecular biology, “omics” technology and advanced microscopy can provide new insight about these mechanisms of interaction between AMF and plants, as well as other microbes. This review mainly discusses the effect of salinity on AMF and plants, and role of AMF in alleviation of salinity stress including insight on methods for AMF identification. The focus remains on latest advancements in mycorrhizal research that can potentially offer an integrative understanding of the role of AMF in salinity tolerance and sustainable crop production.     

AM真菌对盐胁迫下番茄幼苗生理特征及AVP1表达的影响

采用温室盆栽试验研究不同NaCl浓度（0、50 和85 mmol/L）持续胁迫接种摩西球囊霉和地表球囊霉 2种AM真菌对加工番茄耐盐性的影响。结果显示:（1）在0 mmol/L NaCl处理条件下,2种菌的番茄菌根化苗的根系活力、叶片中可溶性糖、可溶性蛋白、根系脯氨酸含量以及超氧化物歧化酶和过氧化物酶活性均高于非菌根植株,且丙二醛含量低于非菌根植株,但差异不显著。（2）在50、85 mmol/L NaCl浓度胁迫下,接种2种菌根真菌可显著提高番茄植株根系活力,促进叶片中可溶性糖、可溶性蛋白及根系脯氨酸含量的积累,显著提高叶片中与抗逆相关的超氧化物歧化酶和过氧化物酶的活性,减少丙二醛在根系中的积累;随着NaCl浓度的增加,效果更为明显。（3）RT-PCR分析显示,AM真菌和盐胁迫共同调控H⁺转运无机焦磷酸酶H⁺- PPase的表达,随NaCl浓度的增加,AVP1基因表达量下降,但菌根化番茄植株的AVP1基因表达量显著高于非菌根植株。研究表明,接种AM真菌后,菌根化植株可通过显著促进幼苗体内渗透调节物质积累和抗氧化酶活性的提高,有效降低体内膜脂过氧化水平,同时过量表达AVP1基因增加了番茄植株中离子向液泡膜的转运,从而缓解盐胁迫对植株的伤害,增强番茄幼苗对盐胁迫的耐性。     

A meta-analysis of arbuscular mycorrhizal effects on plants grown under salt stress

Salt stress limits crop yield and sustainable agriculture in most arid and semiarid regions of the world. Arbuscular mycorrhizal fungi (AMF) are considered bio-ameliorators of soil salinity tolerance in plants. In evaluating AMF as significant predictors of mycorrhizal ecology, precise quantifiable changes in plant biomass and nutrient uptake under salt stress are crucial factors. Therefore, the objective of the present study was to analyze the magnitude of the effects of AMF inoculation on growth and nutrient uptake of plants under salt stress through meta-analyses. For this, data were compared in the context of mycorrhizal host plant species, plant family and functional group, herbaceous vs. woody plants, annual vs. perennial plants, and the level of salinity across 43 studies. Results indicate that, under saline conditions, AMF inoculation significantly increased total, shoot, and root biomass as well as phosphorous (P), nitrogen (N), and potassium (K) uptake. Activities of the antioxidant enzymes superoxide dismutase, catalase, peroxidase, and ascorbate peroxidase also increased significantly in mycorrhizal compared to nonmycorrhizal plants growing under salt stress. In addition, sodium (Na) uptake decreased significantly in mycorrhizal plants, while changes in proline accumulation were not significant. Across most subsets of the data analysis, identities of AMF (Glomus fasciculatum) and host plants (Acacia nilotica, herbs, woody and perennial) were found to be essential in understanding plant responses to salinity stress. For the analyzed dataset, it is concluded that under salt stress, mycorrhizal plants have extensive root traits and mycorrhizal morphological traits which help the uptake of more P and K, together with the enhanced production of antioxidant enzymes resulting in salt stress alleviation and increased plant biomass.     

丛枝菌根真菌对盐胁迫下黄瓜幼苗渗透调节物质含量和抗氧化酶活性的影响

以盐敏感型黄瓜品种‘津春2号’为材料,研究了丛枝菌根真菌(AMF)对盐胁迫下黄瓜幼苗生长及叶片、根系中渗透调节物质含量和抗氧化酶活性的影响.结果表明:(1)在盐胁迫条件下,黄瓜幼苗生长受到明显抑制,其株高、地上部、地下部干鲜重均明显减小,同时体内可溶性蛋白、可溶性糖、脯氨酸和MDA含量,以及O2(÷)产生速率和SOD、POD、CAT活性均比对照显著升高.(2)盐胁迫下接种AMF可显著促进黄瓜植株的生长,进一步提高黄瓜幼苗体内可溶性蛋白、可溶性糖和脯氨酸含量及SOD、POD、CAT活性,而显著降低MDA含量和O2(÷)产生速率.研究表明,AMF可通过显著促进盐胁迫下黄瓜幼苗体内渗透调节物质积累和抗氧化酶活性提高,有效降低体内膜脂过氧化水平,从而缓解盐胁迫对植株的伤害,增强黄瓜幼苗对盐胁迫的耐性.     

The combination of arbuscular mycorrhizal fungi inoculation (Glomus versiforme) and 28‐homobrassinolide spraying intervals improves growth by enhancing photosynthesis,nutrient absorption,and antioxidant system in cucumber (Cucumis sativus L.) under salinity

Salinity is one of the major obstacles in the agriculture industry causing huge losses in productivity. Several strategies such as plant growth regulators with arbuscular mycorrhizal fungi (AMF) have been used to decrease the negative effects of salt stress. In our experiment, 28‐homobrassinolide (HBL) with spraying intervals was combined with AMF (Glomus versiforme) in cucumber cultivars Jinyou 1^# (salt sensitive) and (Changchun mici, in short, CCMC, salt tolerant) under NaCl (100 mmol/L). Studies have documented that the foliar application of HBL and AMF colonization can enhance tolerance to plants under stress conditions. However, the mechanism of the HBL spraying intervals after 15 and 30 days in combination with AMF in cucumber under salt stress is still unknown. Our results revealed that the HBL spraying interval after 15 days in combination with AMF resulted in improved growth, photosynthesis, and decreased sodium toxicity under NaCl. Moreover, the antioxidant enzymes SOD (superoxide dismutase; EC 1.15.1.1) and POD activity (peroxidase; EC 1.11.1.7) showed a gradual increase after every 10 days, while the CAT (catalase; EC 1.11.1.6) increased after 30 days of salt treatments in both cultivars. This research suggests that the enhanced tolerance to salinity was mainly related to elevated levels of antioxidant enzymes and lower uptake of Na⁺, which lowers the risk of ion toxicity and decreases cell membrane damage.     

Characterization of salt-tolerance mechanisms in mycorrhizal (Claroideoglomus etunicatum) halophytic grass,Puccinellia distans

Responses of Puccinellia distans, a halophytic grass to low (50 mM) and high (200 mM) NaCl salinity, were studied in a sand culture experiment without or with inoculation by arbuscular mycorrhizal fungus (AMF), Claroideoglomus etunicatum isolated from its saline habitat. Plant biomass was not influenced by salinity levels, while a tendency to a higher biomass was observed in AMF plants under both control and saline conditions. Leaf photosynthesis increased by both salinity and AMF inoculation. Despite higher transpiration rate, AMF plants had higher water-use efficiency under sever saline conditions. AMF inoculation decreased proline concentration, but increased significantly leaf osmotic potential. Antioxidative enzymes responded differently to the salt and AMF treatments depending on the salt concentration and plant organ. Nonetheless, salt-induced malondialdehyde accumulation in the leaves diminished by AMF colonization. K and Ca contents were not affected by salt, while fungal colonization increased K in the roots and Ca in both leaves and roots. Our results indicated that enhancement of photosynthesis and ion homeostasis is involved in the tolerance of P. distans to both low and high salinity. AMF inoculation increased plants’ tolerance by augmentation of the above mechanisms accompanied by improvement of water relations and protection against oxidative damage in the leaves.