丛枝菌根真菌(AMF)对植物群落调节的研究进展

1 引言 菌根(mycorrhiza)是土壤中的菌根真菌与高等植物营养根系形成的一种共生体,菌根的个主要的类型(即外生菌根Ectomycorrhiza、内生菌根Endomycorrhiza、内外生菌根Ectendomycorrhiza)中,内生性的丛枝状菌根(Vesicular-Arbuscular mycorrhiza,AM)是分布最广泛、最普遍的一类菌根。土壤中的丛枝菌根真菌(Abuscular mycorrhizal fungi, AMF)与高等植物营养根系形成丛枝菌根(abuscular mycorrhiza, AM),能促进宿主对土壤中矿质元素P、NK、Cu、Zn等的吸收,提高宿主根系对根部侵染病菌的抵抗能力和增强植物对干旱、高温、高盐和…     

丛枝菌根（AM）真菌对翅果油树幼苗根系的影响

采用接种和不接种(对照)丛枝菌根的方法对翅果油树幼苗在不同生长时期的形态、生物量、矿质元素含量及抗逆性等指标进行比较分析,以探讨丛枝菌根(AM)改善翅果油树在生态系统中的退化及濒危现状的机理.结果显示:(1)摩西球囊霉(Glomus mosseae)和脆无梗囊霉(Acaulospora delicata)与盆栽翅果油树苗木可形成菌根;菌根化苗木的形态及生物量等指标均与对照差异极显著,且与菌根侵染率呈显著正相关(P<0.05).(2)菌根使苗木根系体积增大、总吸收面积增加、根系的活跃吸收面积显著增加,且根系中活跃吸收面积比与磷、钾元素含量呈显著正相关(P<0.05).(3)苗木生长的盛期和末期,菌根化苗木根系过氧化物酶活性显著高于对照,而在生长后期却显著低于对照呈负相关(P<0.05);菌根化苗木根系多酚氧化酶活性显著高于对照(P<0.05),在苗木生长盛期酶活性最强,但接种不同菌种的苗木间差异不显著.研究表明,与不接种相比,接种菌根增加了翅果油树的根系吸收能力,提高了根系酶体系,有利于植物抵抗各种胁迫,对扩大翅果油树植物的分布区有重要意义.     

纳米氧化锌和接种丛枝菌根真菌对大豆生长及营养吸收的影响

纳米氧化锌是应用最广的人工纳米颗粒(nanoparticles,NPs)之一,具有一定生物毒性。丛枝菌根(arbuscular mycorrhizal,AM)真菌能与陆地上80%以上的高等植物形成丛枝菌根共生体,并能改善宿主植物矿质营养,提高其抗逆性。然而纳米ZnO与丛枝菌根的关系尚不清楚。通过温室沙培盆栽试验,研究了施加不同水平纳米ZnO(0、500、1000、2000、3000 mg/kg)和接种AM真菌Acaulospora mellea对大豆生长及营养状况的影响。结果表明,3000 mg/kg的纳米ZnO显著抑制大豆植株生长,表现出植物毒性,在其他水平时没有显著影响。纳米ZnO在施加水平500、1000 mg/kg时没有抑制AM真菌对大豆根系的侵染,但是高施加水平(2000 mg/kg)时对AM真菌产生毒害,几乎完全抑制大豆根系菌根侵染。接种AM真菌仅在500 mg/kg纳米ZnO时显著促进大豆生长,增加大豆植株对P、K、N的吸收,降低根系Zn含量。纳米ZnO可能会持续释放锌离子,并抑制大豆根系对矿质营养元素的吸收,从而产生生物毒性,而AM真菌与大豆根系的共生可起到有益作用。     

AMF增强枳抗旱性作用机制的初步研究

丛枝菌根真菌(AMF)能增强寄主植物的抗旱性,但相关机制仍不清楚。本研究分析了正常水分和干旱处理对比下AM真菌Diversispora versiformis对盆栽枳(Poncirus trifoliata)生长、光合作用和叶片激素含量的影响。结果表明,6周的干旱处理强烈抑制D.versiformis对枳根系的侵染。干旱处理下,AM真菌接种处理均显著提高了叶、茎和根系生物量,显著增强了叶片光合速率、蒸腾速率和气孔导度,同时显著降低了叶片温度(P0.05)。AM真菌接种也增加了干旱条件下枳叶片内源脱落酸、吲哚乙酸、赤霉素、油菜素内酯、茉莉酸甲酯、玉米素核苷和一氧化氮含量。推测AM真菌通过改变植物内源激素平衡状况,进而诱导相关激素信号传导进程,从而参与菌根的形成、植物生长进程,并增强植物抗旱性。     

丛枝霸王（Zygophyllum dumosum）根际AM真菌生态学研究

AM真菌是一类广泛分布的土壤真菌,与宿主植物形成共生结构后,对于植物生长和植被恢复有多种有益的生理学和生态学作用。1999年11月至2000年10月,通过每月分别从0－10cm和10－20cm土层采集根际土样,对以色列荒漠地区丛枝霸王（Zygophyllum dumosum Boiss）根际AM真菌进行了系统的生态学研究。AM真菌的分布和定殖与季节变化和采样浓度密切相关。菌根真菌的最高定殖率并不伴随有最大的孢子密度,最高的定殖率发生在1999年11月,而最大的孢子密度则出现在2000年9月。10－20cm土层中的菌根真菌定殖率和孢子密度明显高于0－10cm土层。土壤温度与所有菌根结构的定殖率呈正相关,土壤有机质含量与泡囊和丛枝定殖率呈正相关,而土壤总可溶性氮对泡囊和丛枝定殖有显著正效应,对孢子密度有显著负效应。结果建议,孢子密度、泡囊和丛枝定殖程度可作为检测荒漠土壤状况的生态指标。研究应用于我国特别是西部荒漠地区的植被建立和恢复,可望发挥重任作用：（1）AM真菌能与绝大多数高奶系形成共生联合体,促进根系对土壤矿质营养和水分的吸收,提高植物对干旱、高温、盐碱、根部病害等的抗性,提高逆境条件下植物的成活率,深入研究荒漠生态系统中AM真菌动态分布,以及筛选优势AM菌种和人工接种,进行菌根化育苗,为植被建立和恢复提供优质苗木;（2）菌根的丛枝定殖时间短,主要发生在幼根,泡囊定殖时间长,主要发生在老根,而AM真菌的生长发育和繁殖所需的碳水化合物来自植物根系的分泌活动,所以,通过检测不同时期菌根各种结构的定殖程度和孢子的丰富度,可以获得宿主植物根系的生长状况,进而对土壤环境作出科学的评估。     

AM真菌与非菌根植物的相互作用关系

丛枝菌根(AM)真菌作为一类在全球分布广泛的土壤微生物,能够与陆地上大多数的维管植物形成专性共生关系,对于植物营养吸收和生态系统功能具有重要作用.而较少量的维管植物如苋科、黎科、石竹科、十字花科等植物被认为是非菌根植物.目前,对于这些非菌根植物与AM真菌之间的相互作用关系研究少且分散,缺乏系统总结.本文综述了非菌根植物的类型以及低侵染的原因,邻体植物形成的菌丝网络对AM真菌侵染非菌根植物的影响,并探讨AM真菌和非菌根植物之间可能存在的相互作用,以及植物-AM真菌之间的物质交换及可能存在的生态功能,旨在为进一步发挥非菌根植物在脆弱生态系统的功能潜力提供新思路.     

AM真菌氮代谢与运转研究新进展

<正>丛枝菌根(arbuscular mycorrhiza,AM)真菌是一类古老、分布广泛的菌物,能与陆地上80%以上的植物根系建立共生关系,形成共生     

植物中丛枝菌根形成的信号途径研究进展

丛枝菌根(arbuscular mycorrhizal,AM)共生是丛枝菌根真菌与大多数陆地植物的根系之间形成的一种互利共生关系。植物给菌根真菌提供碳水化合物;作为回报,菌根真菌能够增强植物对矿质营养元素(尤其是磷)的吸收。菌根的形成过程是一系列信号交换和转导的结果,具有严格并且一致的顺序。本文以植物中菌根形成的信号途径为主线,对菌根真菌的形成过程和信号转导途径及其方式进行了分析和讨论。高等植物中菌根形成的信号途径与豆科植物的结瘤信号途径部分共享,并且与钙离子信号途径相关,但前者更为广泛。尽管该途径中很多过程目前还不十分清楚,但是相信在不久的将来就可以揭开菌根形成过程中的众多谜团。     

植物中丛枝菌根形成的信号途径研究进展

丛枝菌根(arbuscular mycorrhizal, AM)共生是丛枝菌根真菌与大多数陆地植物的根系之间形成的一种互利共生关系。植物给菌根真菌提供碳水化合物; 作为回报, 菌根真菌能够增强植物对矿质营养元素(尤其是磷)的吸收。菌根的形成过程是一系列信号交换和转导的结果, 具有严格并且一致的顺序。本文以植物中菌根形成的信号途径为主线, 对菌根真菌的形成过程和信号转导途径及其方式进行了分析和讨论。高等植物中菌根形成的信号途径与豆科植物的结瘤信号途径部分共享, 并且与钙离子信号途径相关, 但前者更为广泛。尽管该途径中很多过程目前还不十分清楚, 但是相信在不久的将来就可以揭开菌根形成过程中的众多谜团。     

丛枝菌根对有机污染土壤的修复作用及机理

丛枝菌根(AM)是丛枝菌根真菌(AMF)与植物根系相互作用的互惠共生体,能改良土壤结构,增强植物抗性.自然界中已知的AMF有170多种,分布广泛,且可与大多数植物共生.利用AM修复有机污染土壤正成为一个崭新的研究方向.本文综述了AM对多环芳烃、酞酸脂、石油和农药等一些典型有机污染物污染土壤的修复作用.AM修复有机污染土壤的机理主要包括:AMF代谢有机污染物;AM分泌酶,降解污染物;AM影响根系分泌作用,并促进根际微生物对有机污染物的降解;AMF宿主植物吸收积累污染物.AM修复研究中,高效AMF的筛选、复合菌种效应、土壤老化、AM作用下植物对有机污染物的吸收积累等几方面仍有待于深入研究.     

Localized and non‐localized effects of arbuscular mycorrhizal symbiosis on accumulation of osmolytes and aquaporins and on antioxidant systems in maize plants subjected to total or partial root drying

The arbuscular mycorrhizal (AM) symbiosis alters host plant physiology under drought stress, but no information is available on whether or not the AM affects respond to drought locally or systemically. A split‐root system was used to obtain AM plants with total or only half root system colonized as well as to induce physiological drought affecting the whole plant or non‐physiological drought affecting only the half root system. We analysed the local and/or systemic nature of the AM effects on accumulation of osmoregulatory compounds and aquaporins and on antioxidant systems. Maize plants accumulated proline both, locally in roots affected by drought and systemically when the drought affected the whole root system, being the last effect ampler in AM plants. PIPs (plasma membrane intrinsic proteins) aquaporins were also differently regulated by drought in AM and non‐AM root compartments. When the drought affected only the AM root compartment, the rise of lipid peroxidation was restricted to such compartment. On the contrary, when the drought affected the non‐AM root fraction, the rise of lipid peroxidation was similar in both root compartments. Thus, the benefits of the AM symbiosis not only rely in a lower oxidative stress in the host plant, but it also restricts locally such oxidative stress.     

Azospirillum and arbuscular mycorrhizal colonization enhance rice growth and physiological traits under well-watered and drought conditions

The response of rice plants to inoculation with an arbuscular mycorrhizal (AM) fungus, Azospirillum brasilense, or combination of both microorganisms, was assayed under well-watered or drought stress conditions. Water deficit treatment was imposed by reducing the amount of water added, but AM plants, with a significantly higher biomass, received the same amount of water as non-AM plants, with a poor biomass. Thus, the water stress treatment was more severe for AM plants than for non-AM plants. The results showed that AM colonization significantly enhanced rice growth under both water conditions, although the greatest rice development was reached in plants dually inoculated under well-watered conditions. Water level did not affect the efficiency of photosystem II, but both AM and A. brasilense inoculations increased this value. AM colonization increased stomatal conductance, particularly when associated with A. brasilense, which enhanced this parameter by 80% under drought conditions and by 35% under well-watered conditions as compared to single AM plants. Exposure of AM rice to drought stress decreased the high levels of glutathione that AM plants exhibited under well-watered conditions, while drought had no effect on the ascorbate content. The decrease of glutathione content in AM plants under drought stress conditions led to enhance lipid peroxidation. On the other hand, inoculation with the AM fungus itself increased ascorbate and proline as protective compounds to cope with the harmful effects of water limitation. Inoculation with A. brasilense also enhanced ascorbate accumulation, reaching a similar level as in AM plants. These results showed that, in spite of the fact that drought stress imposed by AM treatments was considerably more severe than non-AM treatments, rice plants benefited not only from the AM symbiosis but also from A. brasilense root colonization, regardless of the watering level. However, the beneficial effects of A. brasilense on most of the physiological and biochemical traits of rice plants were only clearly visible when the plants were mycorrhized. This microbial consortium was effective for rice plants as an acceptable and ecofriendly technology to improve plant performance and development.     

Colonization with arbuscular mycorrhizal fungus affects growth,drought tolerance and expression of stress-responsive genes in <Emphasis Type="Italic">Poncirus trifoliata</Emphasis>

The objective of this study was to investigate the effects of arbuscular mycorrhizal fungus (AMF) inoculation on growth and drought tolerance of Poncirus trifoliata seedlings. The seedlings were inoculated with or without Glomus mosseae before exposure to a short-term (3 days) water depletion, and relevant physiological and biochemical parameters (plant height, chlorophyll content, relative water content, activity of antioxidant enzymes) and expression patterns of several stress-responsive genes were examined. Inoculation with G. mosseae led to growth promotion of the seedlings, as revealed by larger plant height and higher relative water and chlorophyll contents. When subjected to drought treatment, the AMF-inoculated (AM) plants showed better tolerance than the nonmycorrhizal (NAM) plants. Under drought, the AM plants exhibited higher level of proline and activity of two antioxidant enzymes, superoxide dismutase (SOD) and peroxidase (POD). In addition, mRNA abundance of four genes involved in reactive oxygen species homeostasis and oxidative stress battling was higher in the AM plants when compared with the NAM plants. These results indicate that AMF inoculation stimulated growth and enhanced drought tolerance of the seedlings, which may be due to activation of an arsenal of physiological, biochemical and molecular alterations.     

Mycorrhizal symbiosis and response of sorghum plants to combined drought and salinity stresses

Arbuscular mycorrhizal (AM) symbiosis can confer increased host resistance to drought stress, although the effect is unpredictable. Since AM symbiosis also frequently increases host resistance to salinity stress, and since drought and salinity stress are often linked in drying soils, we speculated that the AM influence on plant drought response may be partially the result of AM influence on salinity stress. We tested the hypothesis that AM-induced effects on drought responses would be more pronounced when plants of comparable size are exposed to drought in salinized soils. In two greenhouse experiments, several water relations characteristics were measured in sorghum plants colonized by Glomus intraradices (Gi), Gigaspora margarita (Gm) or a mixture of AM species, during a sustained drought following exposure to salinity treatments (NaCl stress, osmotic stress via concentrated macronutrients, or soil leaching). The presence of excess salt in soils widened the difference in drought responses between AM and nonAM plants in just two instances. Days required for plants to reach stomatal closure were similar for Gi and nonAM plants exposed to drought alone, but with exposure to combined NaCl and drought stress, stomates of Gi plants remained open 17-22% longer than in nonAM plants. Promotion of stomatal conductance by Gm occurred with exposure to NaCl/drought stress but not with drought alone or with soil leaching before drought. In other instances, however, the addition of salt tended to nullify an AM-induced change in drought response. Our findings confirm that AM fungi can alter host response to drought but do not lend much support to the idea that AM-induced salt resistance might help explain why AM plants can be more resilient to drought stress than their nonAM counterparts.     

小麦根系特征对干旱胁迫的响应

干旱胁迫时, 小麦(Triticum aestivum)根系率先产生应激响应, 同时向地上部发出信号, 诱导地上部发生生理反应, 从而提高植株抗旱能力。根系构型包括平面几何性状和立体几何结构(即拓扑构型), 具有遗传稳定性和可塑性。干旱胁迫影响根系理化特性, 如根源化学信号、根系细胞酶类和根系渗透作用的响应。根系通过调整其解剖学结构和水分吸收动力等来适应干旱胁迫。该文从根系构型、理化特性和解剖学结构3个方面, 系统阐述了小麦根系特征对干旱胁迫的响应, 并探讨了其与干旱胁迫的关系和当前研究中存在的问题, 以期为相关研究提供参考。     

小麦根系特征对干旱胁迫的响应

干旱胁迫时, 小麦(Triticum aestivum)根系率先产生应激响应, 同时向地上部发出信号, 诱导地上部发生生理反应, 从而提高植株抗旱能力。根系构型包括平面几何性状和立体几何结构(即拓扑构型), 具有遗传稳定性和可塑性。干旱胁迫影响根系理化特性, 如根源化学信号、根系细胞酶类和根系渗透作用的响应。根系通过调整其解剖学结构和水分吸收动力等来适应干旱胁迫。该文从根系构型、理化特性和解剖学结构3个方面, 系统阐述了小麦根系特征对干旱胁迫的响应, 并探讨了其与干旱胁迫的关系和当前研究中存在的问题, 以期为相关研究提供参考。     

The application of a treated sugar beet waste residue to soil modifies the responses of mycorrhizal and non mycorrhizal lettuce plants to drought stress

Positive effects of sugar beet (SB) application on soil properties and performance of several woody shrub legumes have been described under heavy metal stress and in diverse degraded environments, especially when combined with arbuscular mycorrhiza (AM). However, information on the combined effect of SB amendments and AM symbiosis in horticultural crop plants under drought stress is scarce. Thus, the main objective of this work was to determine if the combination of treated SB waste and AM fungi results in improved drought tolerance of an horticultural food crop such as lettuce and whether or not the effects observed are linked to enhanced antioxidant activities and regulation of two stress-related genes. Lettuce plants inoculated or not with Glomus intraradices and grown on soil amended or not with a treated SB waste were cultivated under well-watered conditions or subjected to drought stress. Plant growth, expression of two drought responsive genes encoding for ??¹-pyrroline-5-carboxylate synthetase and 9-cis-epoxycarotenoid dioxygenase, oxidative damage to lipids and the activity of four antioxidant enzymes were measured. Results showed that the application of treated SB waste resulted negative for the development of AM and nonAM plants (both under well-watered and under drought stress conditions). This effect can not be ascribed to the impairment of specific plant antioxidant defenses. In contrast, a lack of induction of a gene from the ABA biosynthetic pathway was observed in SB-treated plants, which could have contributed to the low performance of these plants. The positive effects of combined application of treated SB waste as amendment and AM fungi have not been shown for a horticultural food crop such as Lactuca sativa. Thus, before starting a program aimed at the utilization of different amendments based on transformed wastes, basic studies on functional and physiological compatibility between the plant and the amendment are necessary.     

Mycorrhizal and non-mycorrhizal Lactuca sativa plants exhibit contrasting responses to exogenous ABA during drought stress and recovery

The arbuscular mycorrhizal (AM) symbiosis enhances plant tolerance to water deficit through the alteration of plant physiology and the expression of plant genes. These changes have been postulated to be caused (among others) by different contents of abscisic acid (ABA) between AM and non-AM plants. However, there are no studies dealing with the effects of exogenous ABA on the expression of stress-related genes and on the physiology of AM plants. The aim of the present study was to evaluate the influence of AM symbiosis and exogenous ABA application on plant development, physiology, and expression of several stress-related genes after both drought and a recovery period. Results show that the application of exogenous ABA had contrasting effects on AM and non-AM plants. Only AM plants fed with exogenous ABA maintained shoot biomass production unaltered by drought stress. The addition of exogenous ABA enhanced considerably the ABA content in shoots of non-AM plants, concomitantly with the expression of the stress marker genes Lsp5cs and Lslea and the gene Lsnced. By contrast, the addition of exogenous ABA decreased the content of ABA in shoots of AM plants and did not produce any further enhancement of the expression of these three genes. AM plants always exhibited higher values of root hydraulic conductivity and reduced transpiration rate under drought stress. From plants subjected to drought, only the AM plants recovered their root hydraulic conductivity completely after the 3 d recovery period. As a whole, the results indicate that AM plants regulate their ABA levels better and faster than non-AM plants, allowing a more adequate balance between leaf transpiration and root water movement during drought and recovery.     

石生南亚毛灰藓在不同温度和干旱条件下的生理生化特性

通过模拟高温和干旱处理,对喀斯特石漠化生境中南亚毛灰藓(Homomallium simlaense(Mitt.)Broth.Mitt)在胁迫条件下生理特征的变化进行了研究。结果表明,南亚毛灰藓在高温和干旱条件下,各项生理指标均与相对含水量呈显著正相关;丙二醛、渗透调节物质和叶绿素含量均随处理时间的增加和含水量的降低而减少,但植株仍保持较高的可溶性糖含量以维持渗透压的平衡。在极端干旱和高温的条件下,南亚毛灰藓可通过降低生理活性,保持一定的可溶性糖含量度过胁迫期,同时丙二醛含量保持最低状态。高温和干旱处理结束后,进行复水处理,植株的渗透调节物质和丙二醛含量显著升高,光合作用迅速恢复。研究结果表明,南亚毛灰藓适应干旱和高温的极端条件可能与丙二醛含量有关,但复水结束后丙二醛含量升高,胁迫反而增强,说明南亚毛灰藓对高温和干旱具有一定耐受性,原因可能与其长期生存于喀斯特的石生环境有关。     

Mycorrhizal impact on drought stress tolerance of rose plants probed by chlorophyll a fluorescence, proline content and visual scoring

Micropropagated rose plants (Rosa hybrida L., cv. New Dawn) were inoculated with the arbuscular mycorrhizal (AM) fungus Glomus intraradices (Schenk and Smith) and subjected to different drought regimens. The dual objectives of these experiments were to investigate the mechanism and the extent to which AM can prevent drought damages and whether physiological analyses reveal enhanced drought tolerance of an economically important plant such as the rose. In a long-term drought experiment with four different water regimens, visual scoring of wilt symptoms affirmed that AM in a selected host–symbiont combination increased plant performance. This effect was mostly expressed if moderate drought stress was constantly applied over a long period. In a short-term experiment in which severe drought stress was implemented and plants were allowed to recover after 4 or 9 days, no visual differences between mycorrhizal and non-mycorrhizal roses were observed. Therefore, the early physiological steps conferring drought tolerance were prone to investigation. Proline content in leaves proved to be an unsuitable marker for AM-induced drought tolerance, whereas analysis of chlorophyll a fluorescence using the JIP test (collecting stress-induced changes of the polyphasic O-J-I-P fluorescence kinetics in a non-destructive tissue screening) was more explanatory. Parameters derived from this test could describe the extent of foliar stress response and help to differentiate physiological mechanisms of stress tolerance. AM led to a more intense electron flow and a higher productive photosynthetic activity at several sites of the photosynthetic electron transport chain. A K step, known as a stress indicator of general character, appeared in the fluorescence transient only in drought-stressed non-mycorrhizal plants; conversely, the data elucidate a stabilising effect of AM on the oxygen-evolving complex at the donor site of photosystem (PS) II and at the electron-transport chain between PS II and PS I. If drought stress intensity was reduced by a prolonged and milder drying phase, these significant tolerance features were less pronounced or missing, indicating a possible threshold level for mycorrhizal tolerance induction.