外生菌根菌与森林树木的相互关系

生态系统的每个过程都伴随着各种微生物的活动,其中最重要的功能群之一是菌根真菌(菌根菌)。一般认为,菌根菌是自然界多数植物生存最基本的组成部分,陆地上约90％以上的高等植物都具有菌根菌。这些菌类的菌丝体与植物根系结合形成菌根,使植物生长成为可能,使不同种类植物的根系联在一起。根据菌根菌入侵植物根系的方式及菌根的形态特征,菌根可分为外生菌根、内生菌根和内外生菌根3组共7种类型。外生菌根主要出现在松科、桦木科、壳斗科等树种的森林生态系统中,在根系表面形成菌丝鞘,部分菌丝进入根系皮层细胞间隙形成哈氏网表面。菌根菌剂在森林经营中得到广泛地应用。外生菌根菌对森林树木的作用可归纳为：1)促进造林或育苗成活与生长;2)提高森林生态系统中植物的多样性、稳定性和生产力;3)对森林生态系统的综合效应,主要表现在增加植物一土壤联结,改善土壤结构,促进土壤微生物,增强植物器官的功能;4)抗拮植物根部病害病原菌等。树木与菌根菌相互关系研究主要包括：1)菌根共生的机理;2)菌根菌在退化森林生态系统恢复与改造中的作用;3)菌根菌的分布格局与森林生态系统服务功能的关系;4)菌根菌对森林生态系统的综合效应,如菌根菌与森林植物群落结构、物种多样性以及森林系统稳定性和生产力的研究。     

菌根和根分泌物在植物抗重金属中的作用

多年来人们对植物抗重金属的研究一直集中在植物体内的代谢调节和控制上。随着近年来对根际环境的研究,人们发现植物体外的根际环境对植物抗重金属有着重要的影响,目前研究主要集中在对菌根真菌和根系分泌物与植物抗重金属的关系,本文就近年来对菌根真菌和根系分泌物在植物抗重金属毒害中发挥的作用及其可能机理的研究状况作一概述,并对该领域的研究和应用前景作出展望。     

外生菌根对干旱胁迫的响应

从外生菌根真菌、外生菌根共生体以及外生菌根的间接作用等方面阐述外生菌根如何抵制干旱胁迫,并对未来我国外生菌根的研究提出了建议。干旱可以抑制外生菌根真菌的生长并降低其群落中真菌的多样性,干旱胁迫下外生菌根真菌子实体可以利用深度30cm以下的土壤水,子实体的表面积和体积比可作为筛选抗旱真菌的一个重要因子;在遭受干旱胁迫时,外生菌根共生体可以发生形态变化来应对干旱,同时增加了植株水分的吸收并改善了植物的光合作用、活性氧以及激素等相关代谢;外生菌根对植物生长的促进作用、增加土壤碳汇以及对其他根际微生物生长的促进作用等对宿主植物应对干旱胁迫有利。未来我国外生菌根研究应加强对干旱区优良菌-树组合的筛选工作,同时加大对乡土外生菌根真菌资源的调查力度,未来研究应重点向分子生物学领域推进。     

刺槐林下菌根真菌资源的调查、分离及回接

植物与微生物的共生体在长期协同进化的过程中形成了对各种胁迫的适应性。在北方干旱、半干旱的逆境条件下植物生长除受水分制约外 ,氮、磷元素的供应也是重要的限制性因子。固氮植物可以通过根瘤进行固氮作用弥补氮源不足 ,而菌根能够提高宿主植物对磷元素的吸收。研究表明 ,植物感染菌根后可以提高宿主的抗旱性、抗盐碱性、抗重金属毒害、抗极端湿度、抗极端 ｐＨ等的能力[1]。近年来 ,菌根技术被应用于营林实践中 ,在进行某些树种造林时 ,利用人工接种外生菌根真菌的方法 ,已取得初步的成功[2 ]。在我国广大农村 ,薪材在能源消费中占有重…     

外生菌根缓解植物酸雨胁迫的机理研究进展

森林作为陆地生态系统的主体,是酸雨污染的主要受体,酸雨对生态系统产生着巨大的影响。菌根是菌根真菌与植物营养根的共生体。外生菌根真菌与宿主植物间互惠互利,在森林生态系统中,外生菌根在维持生态系统的养分平衡和改善树木营养等方面有重要的作用。本文综述了国内外关于菌根和酸雨关系的研究,酸雨能抑制外生菌根的形成,降低其活力;但另一方面,外生菌根能够缓解酸雨造成的植物危害,提高植株对酸雨的耐受力。外生菌根主要通过以下几方面缓解酸雨胁迫:(1)菌根形态结构的物理屏蔽作用;(2)增加养分吸收,增加御酸能力;(3)增强酶活性,提高植物生存能力;(4)产生有机酸或其他物质。     

不同生境外生菌根真菌对铝胁迫的响应

以南北方不同生境下的10株外生菌根真菌为研究对象,采用液体培养的方法,研究了铝对不同菌根真菌的生物量、有机酸分泌及养分含量的影响,以期筛选出抗铝性强的优良菌株,并探讨其抗铝机理。结果表明:外生菌根真菌Sl 08抗铝性最强;Pt 715、Ld 03、Bo 11、Sl 01、Bo 15也具有不同程度的耐铝性;Sl 14、Gc 99、Cg 04抗铝性较差;Sg 11抗铝性最差。来自南方酸性森林土壤的菌株总体抗铝性强于来自北方石灰性土壤的菌株,这表明外生菌根真菌的铝耐受能力与其原始生境有着密切的联系。外生菌根真菌能分泌多种有机酸,且不同菌株分泌的有机酸种类不同。其中,受铝胁迫分泌量增加最多的是草酸。研究中,铝胁迫能增加大多数铝抗性菌株的草酸分泌量,其中铝抗性最强的Sl 08表现最为明显。但铝胁迫并没有促进具备一定铝抗性的Bo 11和Sl 01草酸的分泌量,同时在铝敏感的菌株中均观察到了草酸分泌量的增加。这表明分泌草酸可能并不是外生菌根真菌抵抗铝毒的唯一途径。对各菌株铝胁迫下对氮,磷及钾的吸收研究表明,除铝敏感菌株Sl 14外,铝胁迫均能促进各供试菌株对氮,磷或钾的吸收。综上,在一定铝浓度下,一些外生菌根真菌可通过增加草酸分泌来抵御铝毒。此外,铝胁迫下外生菌根真菌还可通过调控氮、磷、钾等营养元素的吸收来抵抗铝毒,即通过增加对营养元素的吸收来增强其在铝胁迫下的生存能力,这可能是其抵御铝胁迫的应激反应之一。     

接种彩色豆马勃对模拟酸沉降下马尾松幼苗生物量的影响

外生菌根能够提高宿主植物对外界环境胁迫的抵抗力,促进植物的生长,本文试图揭示外生菌根对酸雨胁迫下马尾松生长的保护作用。本研究采用盆栽试验,共设置四个处理：酸雨对照处理（Control check (CK), 约pH值5.5）不接种,酸雨对照处理接种,酸雨pH值3.5处理不接种,酸雨pH值3.5处理接种。pH值3.5的酸雨处理降低马尾松的生物量,在试验前期降低根冠比,试验中后期则提高根冠比,在试验初期增加了叶面积,但中后期显著降低了叶面积。接种外生菌根菌有利于马尾松幼苗的生长,pH值3.5处理下接种外生菌根菌能提高马尾松幼苗的生物量,外生菌根菌对生物量分配和叶面积的影响与酸雨胁迫的影响是相反的,即外生菌根菌抵消了酸雨胁迫对马尾松的影响。     

杜鹃花科植物菌根的研究进展

杜鹃花科Ericaceae植物可与土壤真菌形成杜鹃花类菌根ericoid mycorrhizas（ERM）共生体,且广泛分布于全球不同的陆地生态系统,特别是在贫瘠、酸性等严酷的环境中占优势。杜鹃花科植物菌根类型多样,绝大多数宿主具有ERM,还有少量宿主具有其他类型的菌根结构,且常与暗隔内生菌（dark septate endophyte,DSE）并存;ERM的宿主植物除已知的杜鹃花科外,岩梅科Diapensiaceae植物也具有ERM结构;ERM真菌以子囊菌和担子菌为主,主要来自柔膜菌目Helotiales和蜡壳耳目Sebacinales;与杜鹃花科宿主形成ERM的真菌也常与壳斗科Fagaceae、松科Pinaceae等宿主植物形成外生菌根（ectomycorrhiza,ECM）结构;ERM对宿主植物在营养吸收、忍耐贫瘠环境、抵抗重金属污染等能力方面都有积极的促进作用,对环境变化的响应是多样的,生境和季节的变化对ERMF群落的组成和分布有着显著影响,资源比率变化可能改变ERM宿主与其他菌根或非菌根植物之间的竞争关系。本文回顾了近40多年来国内外有关ERM的研究进展,还对ERM研究的前景进行了展望,以期在理论和实践中对杜鹃花科及其菌根的研究能取得更丰硕的成果。     

川西亚高山岷江冷杉外生菌根形态随海拔梯度的分化

外生菌根真菌是一类有助于植物生长及应对环境变化的重要共生真菌类群。深入认识外生菌根特征有助于理解植物对环境变化的响应规律。本研究在四川省卧龙自然保护区巴郎山脉设置海拔2850、3000、3194、3413和3593 m 5个样地,利用中心象限法进行土块样品(10 cm×10 cm×10 cm)采集,通过分析各海拔每土块中岷江冷杉外生菌根形态个数、形态多样性及养分获取类型等指标,研究岷江冷杉外生菌根形态特性随海拔的变异特征及土壤环境因子的作用。结果表明: 1) 菌根形态为橘黄色或橘色、单轴羽状分支或不分支、菌套表面光滑、无或少见外延物、根尖直筒型或膨大的菌根类型是卧龙自然保护区岷江冷杉外生菌根的主要形态类型。该形态类型在研究区内的侵染率最高,为12.4%;2) 随着海拔的升高,岷江冷杉外生菌根形态多样性下降,每土块内的形态个数减少;3) 单因素方差分析结果表明,不同海拔上无外延菌丝或极少外延菌丝的紧密接触类型(CE)和短距离养分获取类型(SDE)外生菌根侵染率有显著差异,CE类型外生菌根侵染率随着海拔的升高而增加;4) 土壤环境因子对岷江冷杉外生菌根形态在海拔上的变异有一定的影响,冗余分析(RDA)表明,土壤全氮含量、全磷含量、土壤温度、土壤含水率、pH值及酸性磷酸酶活性对岷江冷杉外生菌根形态在海拔上的变异有显著影响,其中土壤全氮含量和土壤温度的影响最大,分别解释了岷江冷杉形态特征变异的5.4%和4.9%。本研究揭示了岷江冷杉外生菌根形态形成在海拔上的差异,为进一步研究亚高山针叶树种菌根策略对环境变化的响应机制提供了科学依据。     

外生菌根菌的分子生态学：分子标记基因、遗传个体及生态意义

外生菌根真菌与很多植物形成互利共生关系,在营养物质交换和碳循环等方面起着关键性的作用,是森林生态系统的重要组成部分。近期生物技术的发展使得人们对外生菌根菌的群体遗传学和分子生态学有了更加深入的认识。本文介绍了一些常用的鉴定外生菌根菌的分子标记,并对每种分子标记的特点及其适用范围进行了讨论。文中总结了几种常用的鉴定未知外生菌根菌的方法,指出了一些在研究外生菌根菌过程中需要克服的内在困难,其中之一就是很多外生菌根菌不可以人工培养,所以人们缺少对其地下部分分布规律和动态变化的了解。在寄主专一性、物种多样性和丰富度、遗传个体大小、繁殖方式等方面,近期对外生菌根菌的分子生物学研究已经获得了很多重要的结果。作者讨论了这些研究成果对于今后开展外生菌根菌研究的重要意义以及在森林生态系统保育方面的潜在应用价值。     

Mechanisms of metal resistance in plants: aluminum and heavy metals

Plants have evolved sophisticated mechanisms to deal with toxic levels of metals in the soil. In this paper, an overview of recent progress with regards to understanding fundamental molecular and physiological mechanisms underlying plant resistance to both aluminum (Al) and heavy metals is presented. The discussion of plant Al resistance will focus on recent advances in our understanding of a mechanism based on Al exclusion from the root apex, which is facilitated by Al-activated exudation of organic acid anions. The consideration of heavy metal resistance will focus on research into a metal hyperaccumulating plant species, the Zn/Cd hyperaccumulator, Thlaspi caerulescens, as an example for plant heavy metal research. Based on the specific cases considered in this paper, it appears that quite different strategies are used for Al and heavy metal resistance. For Al, our current understanding of a resistance mechanism based on excluding soil-borne Al from the root apex is presented. For heavy metals, a totally different strategy based on extreme tolerance and metal hyperaccumulation is described for a hyperaccumulator plant species that has evolved on naturally metalliferous soils. The reason these two strategies are the focus of this paper is that, currently, they are the best understood mechanisms of metal resistance in terrestrial plants. However, it is likely that other mechanisms of Al and/or heavy metal resistance are also operating in certain plant species, and there may be common features shared for dealing with Al and heavy resistance. Future research may uncover a number of novel metal resistance mechanisms in plants. Certainly the complex genetics of Al resistance in some crop plant species, such as rice and maize, suggests that a number of presently unidentified mechanisms are part of an overall strategy of metal resistance in crop plants.     

Physiological Aspects of Cadmium and Lead Toxic Effects on Higher Plants

Using the examples of cadmium and lead, the review considers the various toxic effects exerted by these heavy metals. Putative specific and nonspecific mechanisms of the toxic effects of the heavy metals and plant responses are discussed together with the issue of Cd and Pb accumulation in various plant organelles, cells, tissues, and organs. The basic mechanisms providing for plant resistance to excess Cd and Pb are elucidated. These data are used to schematically outline the changes in plant metabolism produced by these heavy metals.     

外生菌根对植物非生物胁迫的缓解作用及作用机理研究进展

外生菌根在林木共生系统中占据重要的地位,具有提高宿主植物抗逆性的作用。在菌根学领域中,外生菌根与非生物胁迫互作的研究要远少于其他类型的菌根,尤其是缺乏综合、全面的总结性评述。文中总结了近5年来的相关研究,阐述了非生物胁迫(干旱、寒冷、高温、盐碱、重金属和有毒物质)下菌根共生体与植物抗逆性的关系,其他因子与外生菌根协同提高植物对非生物胁迫的耐受能力,以及外生菌根缓解非生物胁迫的生理机制和基因调节机制。本文结合了污染土壤修复、功能蛋白表达及微生物生态系统3个方面的研究,分析了当前研究的热点问题与存在的不足,展望了今后的研究方向,以期为林木生态恢复和菌根学研究的扩充提供可借鉴的思路。     

Nitrogen fixation associated with Suillus tomentosus tuberculate ectomycorrhizae on Pinus contorta var. latifolia

BACKGROUND AND AIMS: Tuberculate ectomycorrhizae are a unique form of ectomycorrhiza where densely packed clusters of mycorrhizal root tips are enveloped by a thick hyphal sheath to form a tubercle. The functional significance of such a unique structure has not previously been established. The purpose of the present study was to investigate and measure the potential nitrogenase activity associated with Suillus tomentosus/Pinus contorta tuberculate ectomycorrhizae in two stand ages, young and old, and across a range of nitrogen-poor soil conditions. METHODS: Short roots were compared with other mycorrhizae and non-mycorrhizal secondary roots using tuberculate ectomycorrhizae. Assessment of nitrogenase activity was determined and quantitative measurements were taken on tuberculate ectomycorrhizae in situ in a variety of different circumstances, by using an adaptation of the acetylene reduction assay. KEY RESULTS: Significant nitrogenase activity was measured associated with S. tomentosus/P. contorta tuberculate ectomycorrhizae whereas no nitrogenase activity was measured with non-tuberculate mycorrhizae or secondary roots without mycorrhizae. Average nitrogenase activity ranged from undetectable to 5696.7 nmol C2H4 g(-1) tubercle 24 h(-1). Maximum nitrogenase activity was 25,098.8 nmol C2H4 g(-1) tubercle 24 h(-1). Nitrogenase activity was significantly higher in young stands than in old stands of P. contorta. Season or some covariate also seemed to affect nitrogenase activity and there was suggestion of a site effect. CONCLUSIONS: Suillus tomentosus/P. contorta tuberculate ectomycorrhizae are sites of significant nitrogenase activity. The nitrogenase activity measured could be an important contribution to the nitrogen budget of P. contorta stands. Season and stand age affect levels of nitrogenase activity.     

Nutrient amounts of ectomycorrhizae analysed by EDX using ESEM and ICP

Energy-dispersive X-ray (EDX) analysis coupled with an environmental scanning electron microscope (ESEM) was tested as an alternative to the inductively coupled argon plasma (ICP) spectrometer method for nutrient analyses of ectomycorrhizae. The results of EDX–ESEM and ICP were compared for 12 ectomycorrhizal morphotypes collected in beech and Scots pine forests in northern Brandenburg. The amounts of Al, Ca, Mg and S analysed in the outer hyphal layers of the sheath with the EDX–ESEM technique correlated well with the amounts of these elements in the whole mycorrhiza as assessed by ICP. For the elements P and K, no such correlation existed, indicating an uneven distribution of these elements in the ectomycorrhiza. It is concluded that the EDX–ESEM technique could be a useful and reliable tool for the analysis of nutrient elements in ectomycorrhizae, especially for studies focussing on small-scale soil heterogeneity or on infrequent morphotypes.     

锌胁迫对彩色豆马勃体内锌的分布和活性形态及其抗氧化系统的影响

彩色豆马勃Pisolithus tinctorius可广泛与林木建立外生菌根,其共生可促进植物生长,提高植物对重金属的耐受性。然而,关于彩色豆马勃对重金属锌(Zn)胁迫的生理响应还不完全清楚。本研究在纯培养条件下,测定了Zn胁迫对彩色豆马勃生物量、Zn的活性形态和亚细胞分布,超氧化物歧化酶(SOD)、过氧化氢酶(CAT)和过氧化物酶(POD)活性,谷胱甘肽(GSH)、抗坏血酸(AsA)和黑色素含量的影响。结果表明,Zn胁迫抑制了彩色豆马勃的生长,在浓度为563 μmol/L时,达到半数最大抑制浓度(IC₅₀),在浓度为600 μmol/L时,生物量降低42.6%。Zn在彩色豆马勃中以活性较低的草酸盐、难溶于水的磷酸盐、果胶酸盐以及与蛋白质呈结合态或吸着态的形式为主。Zn胁迫显著提高了彩色豆马勃菌丝中CAT、POD和SOD活性,增加了GSH和黑色素含量,但对AsA的含量没有显著影响。本研究表明,彩色豆马勃主要通过调节Zn的分布及抗氧化系统,增加黑色素含量,增强对Zn的耐受性,为外生菌根真菌修复重金属污染土壤提供了理论支持。     

Contrasting the Pb (II) and Cd (II) tolerance of Enterobacter sp. via its cellular stress responses

Successful application of microorganisms to heavy metal remediation depends on their resistance to toxic metals. This study contrasted the differences of tolerant mechanisms between Pb²⁺ and Cd²⁺ in Enterobacter sp. Microbial respiration and production of formic acid showed that Enterobacter sp. had a higher tolerant concentration of Pb (>1000 mg l⁻¹) than Cd (about 200 mg l⁻¹). Additionally, SEM confirmed that most of Pb and Cd nanoparticles (NPs) were adsorbed onto cell membrane. The Cd stress, even at low concentration (50 mg l⁻¹), significantly enlarged the sizes of cells. The cellular size raised from 0.4 × 1.0 to 0.9 × 1.6 μm on average, inducing a platelet-like shape. In contrast, Pb cations did not stimulate such enlargement even up to 1000 mg l⁻¹. Moreover, Cd NPs were adsorbed homogeneously by almost all the bacterial cells under TEM. However, only a few cells work as ‘hot spots’ on the sorption of Pb NPs. The heterogeneous sorption might result from a ‘self-sacrifice’ mechanism, i.e., some cells at a special life stage contributed mostly to Pb sorption. This mechanism, together with the lower mobility of Pb cations, caused higher microbial tolerance and removal efficiency towards Pb²⁺. This study sheds evident contrasts of bacterial resistance to the two most common heavy metals.     

Ectomycorrhizae of Tomentella albomarginata (Thelephoraceae) on Scots pine

 The ectomycorrhizae of Tomentella albomarginata are comprehensively described and compared to ectomycorrhizae of other Tomentella species and to ectomycorrhizae of some members of Thelephoraceae ss. Stalpers and Bankeraceae ss. Stalpers. The ectomycorrhizae of T. albomarginata are characterized by a hyphal net lying on a pseudoparenchymatous mantle surface, by tubular outgrowths of irregularly angular mantle cells, and by clamp-bearing emanating hyphae. Accepted: 15 July 1995