丛枝菌根真菌提高盐胁迫植物抗氧化机制的研究进展

土地盐渍化是在自然环境和人为活动的双重作用下形成的全球性的重要生态问题,其会对植物造成渗透失衡、离子胁迫、氧化损伤等危害,导致植物生长缓慢、生物量减少甚至是绝产。丛枝菌根真菌(AMF)是一种普遍存在于土壤中的有益微生物,能够与大多数植物根系形成共生关系,其共生关系在多种逆境生态系统中均具有重要生态意义。AMF-植物共生体具有高效抗氧化系统,能够提高植物在盐胁迫下的抗氧化反应进而增强耐盐性。本文从氧化损伤、渗透调节、抗氧化机制和生物活性分子等角度,系统地阐述了丛枝菌根真菌提高植物抗氧化机制的研究进展,并提出了研究展望,以期为利用菌根生物技术提高植物耐盐性提供理论参考。     

Benefit, cost and water-use efficiency of arbuscular mycorrhizal durum wheat grown under drought stress

 Arbuscular mycorrhizal fungi (AMF) living symbiotically with host plants enhance plant growth by improving the acquisition of mineral nutrients and water relations. This study determined the effects of AMF inoculation on growth, benefit/cost and water-use efficiency (grams dry matter produced per kilogram water evapotranspired) in two durum wheat genotypes (drought sensitive and drought tolerant) under water-stressed and well-watered conditions. Plants were grown in a low-P silty clay (Typic Xerochrept) soil mix in a greenhouse. Shoot and root dry matter (DM) and root AMF colonization were higher for well-watered than for water-stressed plants. The mycorrhizal plants were more water-use efficient than nonmycorrhizal plants. Shoot DM differences between mycorrhizal and nonmycorrhizal plants represent the benefit derived by plants from AMF-root associations. Shoot DM differences between mycorrhizal and nonmycorrhizal plants under similar conditions of water treatment represent the cost to the plant of AMF-root associations. Values of benefit/cost for AMF-root associations were highest when plants were water-stressed and decreased under well-watered conditions. Genotypic differences in calculated costs and benefits were pronounced. Benefit/cost analysis may be helpful in evaluating host plant genotypes in order to optimize efficiencies of AMF symbiosis under different environmental conditions. Accepted: 4 April 1998     

Influence of arbuscular mycorrhizae on photosynthesis and water status of maize plants under salt stress

The influence of arbuscular mycorrhizal (AM) fungus Glomus mosseae on characteristics of the growth, water status, chlorophyll concentration, gas exchange, and chlorophyll fluorescence of maize plants under salt stress was studied in the greenhouse. Maize plants were grown in sand and soil mixture with five NaCl levels (0, 0.5, 1.0, 1.5, and 2.0 g/kg dry substrate) for 55 days, following 15 days of non-saline pretreatment. Under salt stress, mycorrhizal maize plants had higher dry weight of shoot and root, higher relative chlorophyll content, better water status (decreased water saturation deficit, increased water use efficiency, and relative water content), higher gas exchange capacity (increased photosynthetic rate, stomatal conductance and transpiration rate, and decreased intercellular CO(2) concentration), higher non-photochemistry efficiency [increased non-photochemical quenching values (NPQ)], and higher photochemistry efficiency [increased the maximum quantum yield in the dark-adapted state (Fv/Fm), the maximum quantum yield in the light-adapted sate (Fv'/Fm'), the actual quantum yield in the light-adapted steady state (varphiPSII) and the photochemical quenching values (qP)], compared with non-mycorrhizal maize plants. In addition, AM symbiosis could trigger the regulation of the energy biturcation between photochemical and non-photochemical events reflected in the deexcitation rate constants (kN, kN', kP, and kP'). All the results show that G. mosseae alleviates the deleterious effect of salt stress on plant growth, through improving plant water status, chlorophyll concentration, and photosynthetic capacity, while the influence of AM symbiosis on photosynthetic capacity of maize plants can be indirectly affected by soil salinity and mycorrhizae-mediated enhancement of water status, but not by the mycorrhizae-mediated enhancement of chlorophyll concentration and plant biomass.     

长期定位施肥土壤中AM真菌对寄主植物的侵染状况

分别于2003年9月和2004年5月在莱阳农学院长期(26年)定位施肥试验田采集玉米和小麦根系,以测定不同施肥处理对丛枝菌根(AM)真菌侵染状况的影响。结果表明长期定位施氮(N)肥显著降低了AM真菌对寄主植物的侵染率(MCP)、丛枝着生率(ACP)、单位根长泡囊数(NV)和侵入点数(NE)。高N处理的小麦MCP低于低N处理,高N处理的玉米ACP低于低N处理。长期定位施有机肥,尤其是高有机肥处理显著降低了MCP、ACP和NE。有机肥和N肥配施也降低了MCP、ACP、NV和NE,以高有机肥和高N肥配施处理的降低效应最大。除小麦根系NV外,长期定位NPK配合施用降低了玉米和小麦的MCP、ACP、NV和NE。其中,以NP处理的玉米ACP、小麦的MCP和ACP最低。另外,低有机肥与N肥配施和NPK配施处理条件下玉米MCP高于小麦。结论认为不同施肥体制对作物菌根生长发育及其结构具有不同影响,而且有机肥和N肥在影响AM真菌侵染方面存在互作。     

The differential behavior of arbuscular mycorrhizal fungi in interaction with Astragalus sinicus L. under salt stress

Three arbuscular mycorrhizal (AM) fungi (Glomus mosseae, Glomus claroideum, and Glomus intraradices) were compared for their root colonizing ability and activity in the root of Astragalus sinicus L. under salt-stressed soil conditions. Mycorrhizal formation, activity of fungal succinate dehydrogenase, and alkaline phosphatase, as well as plant biomass, were evaluated after 7 weeks of plant growth. Increasing the concentration of NaCl in soil generally decreased the dry weight of shoots and roots. Inoculation with AM fungi significantly alleviated inhibitory effect of salt stress. G. intraradices was the most efficient AM fungus compared with the other two fungi in terms of root colonization and enzyme activity. Nested PCR revealed that in root system of plants inoculated with a mix of the three AM fungi and grown under salt stress, the majority of mycorrhizal root fragments were colonized by one or two AM fungi, and some roots were colonized by all the three. Compared to inoculation alone, the frequency of G. mosseae in roots increased in the presence of the other two fungal species and highest level of NaCl, suggesting a synergistic interaction between these fungi under salt stress.     

Comparing symbiotic performance and physiological responses of two soybean cultivars to arbuscular mycorrhizal fungi under salt stress

The presented experiments evaluated the symbiotic performance of soybean genotypes with contrasting salt stress tolerance to arbuscular mycorrhizal fungi (AMF) inoculation. In addition, the physiological stress tolerance mechanisms in plants derived from mutualistic interactions between AMF and the host plants were evaluated. Plant growth, nodulation, nitrogenase activity and levels of endogenous growth hormones, such as indole acetic acid and indole butyric acid, of salt-tolerant and salt-sensitive soybean genotypes significantly decreased at 200 mM NaCl. The inoculation of soybean with AMF improved the symbiotic performance of both soybean genotypes by improving nodule formation, leghemoglobin content, nitrogenase activity and auxin synthesis. AMF colonization also protected soybean genotypes from salt-induced membrane damage and reduced the production of hydrogen peroxide, subsequently reducing the production of TBARS and reducing lipid peroxidation. In conclusion, the results of the present investigation indicate that AMF improve the symbiotic performance of soybean genotypes regardless of their salt stress tolerance ability by mitigating the negative effect of salt stress and stimulating endogenous level of auxins that contribute to an improved root system and nutrient acquisition under salt stress.     

The effects of calcium sulphate on growth,membrane stability and nutrient uptake of tomato plants grown under salt stress

A pot experiment was carried out with tomato (Lycopersicon esculentum Mill.) cv. “Target F1” in a mixture of peat, perlite, and sand (1:1:1) to investigate the effects of supplementary calcium sulphate on plants grown at high NaCl concentration (75 mM). The treatments were: (i) control (C), nutrient solution alone; (ii) salt treatment (C + S), 75 mM NaCl; (iii) salt plus calcium treatment 1 (C + S + Ca1), 75 mM NaCl plus additional mixture of 2.5 mM CaSO₄ in nutrient solution; (iv) salt plus calcium treatment 2 (C + S + Ca2), 75 mM NaCl plus additional mixture of 5 mM CaSO₄ in nutrient solution. The plants grown under salt stress produced low dry matter, fruit weight, and relative water content than those grown in standard nutrient solution. Supplemental calcium sulphate added to nutrient solution containing salt significantly improved growth and physiological variables affected by salt stress (e.g. plant growth, fruit yield, and membrane permeability) and also increased leaf K⁺, Ca²⁺, and N in tomato plants. The effects of supplemental CaSO₄ in maintaining membrane permeability, increasing concentrations of Ca²⁺, N, and K⁺ and reducing concentration of Na⁺ (because of cation competition in root zone) in leaves could offer an economical and simple solution to tomato crop production problems caused by high salinity.     

Contribution of six arbuscular mycorrhizal fungal isolates to water uptake by Lactuca sativa plants under drought stress

It is currently accepted that, along with nutrients, arbuscular mycorrhizal (AM) fungi also transport water to their host plant. However, the quantity of water supplied and its significance for plant water relations remain controversial. The objective of this work was to evaluate and compare the ability of six AM fungi to alter rates of root water uptake under drought stress conditions. Soil drying rates of uninoculated control plants of comparable size and nutritional status and mycorrhizal plants were recorded daily. Lactuca sativa plants colonized by Glomus coronatum , G. intraradices , G. claroideum and G. mosseae depleted soil water to a higher extent than comparably sized uninoculated control plants or plants colonized by G. constrictum or G. geosporum . The differences ranged from 0.6% volumetric soil moisture for G. mosseae -colonized plants to 0.95% volumetric soil moisture for G. intraradices -colonized plants. These differences in soil moisture were equivalent to 3–4.75 ml plant⁻¹ day⁻¹, respectively, and could not be ascribed to differences in plant size, but to the activity of AM fungi. The AM fungi tested in this study differed in their effectiveness to enhance plant water uptake from soil. This ability seems to be related to the amount of external mycelium produced by each AM fungus and to the frequency of root colonization in terms of live and active fungal structures.     

Alleviation of salt stress in citrus seedlings inoculated with arbuscular mycorrhizal fungi depends on the rootstock salt tolerance

Seedlings of Cleopatra mandarin (Citrus reshni Hort. ex Tan.) and Alemow (Citrus macrophylla Wester) were inoculated with a mixture of AM fungi (Rhizophagus irregularis and Funneliformis mosseae) (+AM), or left non-inoculated (−AM). From forty-five days after fungal inoculation onwards, half of +AM or −AM plants were irrigated with nutrient solution containing 50 mM NaCl. Three months later, AM significantly increased plant growth in both Cleopatra mandarin and Alemow rootstocks. Plant growth was higher in salinized +AM plants than in non-salinized −AM plants, demonstrating that AM compensates the growth limitations imposed by salinity. Whereas AM-inoculated Cleopatra mandarin seedlings had a very good response under saline treatment, inoculation in Alemow did not alleviate the negative effect of salinity. The beneficial effect of mycorrhization is unrelated with protection against the uptake of Na or Cl and the effect of AM on these ions did not explain the different response of rootstocks. This response was related with the nutritional status since our findings confirm that AM fungi can alter host responses to salinity stress, improving more the P, K, Fe and Cu plant nutrition in Cleopatra mandarin than in Alemow plants. AM inoculation under saline treatments also increased root Mg concentration but it was higher in Cleopatra mandarin than in Alemow. This could explain why AM fungus did not completely recovered chlorophyll concentrations in Alemow and consequently it had lower photosynthesis rate than control plants. AM fungi play an essential role in citrus rootstock growth and biomass production although the intensity of this response depends on the rootstock salinity tolerance.     

Arbuscular mycorrhizal symbiosis influences strigolactone production under salinity and alleviates salt stress in lettuce plants

Arbuscular mycorrhizal (AM) symbiosis can alleviate salt stress in plants. However the intimate mechanisms involved, as well as the effect of salinity on the production of signalling molecules associated to the host plant-AM fungus interaction remains largely unknown. In the present work, we have investigated the effects of salinity on lettuce plant performance and production of strigolactones, and assessed its influence on mycorrhizal root colonization. Three different salt concentrations were applied to mycorrhizal and non-mycorrhizal plants, and their effects, over time, analyzed. Plant biomass, stomatal conductance, efficiency of photosystem II, as well as ABA content and strigolactone production were assessed. The expression of ABA biosynthesis genes was also analyzed.     

Priming of seeds with NaCl induces physiological changes in tomato plants grown under salt stress

The effects of seed priming with 6 M NaCl solution have been investigated with respect to growth and physiological responses of tomato plants ( Lycopersicon esculentum Mill. cv. Pera) exposed to 70 and 140 m M NaCl nutrient solutions from 11 to 60 days after sowing. Tomato seedlings from primed seeds emerged earlier than from non-primed seeds. At 70 m M , a lower shoot and root dry weight reduction was found in plants from primed seeds at the different harvests (30, 45 and 60 days after sowing), while at 140 m M the positive effect of seed priming was only shown in roots. Significant changes in Na⁺ and CI⁻ accumulation with seed priming were only found in roots at 60 days after sowing, with ion accumulation in roots being higher in plants grown at 70 and 140 m M from primed seeds. In leaves of salt-treated plants, significant increases in sugars and organic acids with seed priming were found from 30 days after sowing, and these increases were higher at longer treatment times. In roots, however, only the organic acids tended to increase in plants from primed seeds, although they increased less than in leaves, especially at 60 days after sowing. These results support the hypothesis that priming of seeds with NaCl induces physiological changes in the plants, changes which are shown more clearly at advanced growth stages.     

盐胁迫下外源褪黑素和从枝菌根真菌对月季幼苗生理特性及光合荧光参数的影响

本试验旨在探究100mol/L盐胁迫下根际施用褪黑素(MT)、接种近明球囊霉属AMF幼套近明球囊霉(Claroideoglomus etunicatum)及其复合处理对月季幼苗生长、叶绿素荧光参数、激素代谢及抗氧化系统的影响,以探明两者缓解月季盐胁迫的机制。结果发现,盐胁迫下月季幼苗生长受到抑制,株高、茎粗以及生物量等显著下降;施用MT可以促进AMF侵染,提高侵染率、丛枝着生率、泡囊数和侵入点数。100mol/L盐处理下,与对照(CK)处理相比,AMF+MT处理的叶绿素总量、叶绿素a/b分别增加46.2%和67.2%;叶绿素荧光参数中PSⅡ最大光化学效率(Fv/Fm)、PSⅡ潜在活性(Fv/Fo)、PSⅡ实际光化学效率(φPSⅡ)、PSII有效光化学量子效率(Fv’/ Fm’)、光化学猝灭系数(qP)分别增加4.9%、51.0%、175.0%、168.7%和92.5%,NPQ的下降幅度为42.7%;此外,盐胁迫下,月季叶片中玉米素核苷(ZR)、赤霉素(GA)、生长素(IAA)含量下降,而脱落酸含量(ABA)增加,AMF+MT处理后ZR、GA、IAA分别增加146.9%、116.9%、35.7%,ABA下降21.1%;同时AMF+MT处理能够激活抗氧化酶SOD、CAT活性,降低超氧阴离子(O2-)产生速率和H2O2累积。结论认为,接种AMF、添加MT或者AMF+MT处理均可以提高叶绿素含量,保护叶绿素荧光系统,维持植物内源激素的平衡,激活SOD、CAT等抗氧化酶活性以及降低脂质过氧化和H2O2累积,以减轻盐胁迫对月季幼苗的伤害,促进月季生长,其中以AMF+MT处理下月季幼苗的抗盐性效果更佳。     

Effects of arbuscular mycorrhizal fungi on photosystem II activity of three pistachio rootstocks under salt stress as probed by the OJIP-test

We applied chlorophyll a fluorescence as a biomarker to assess the growth response and PSII behavior and performance of three pistachio (Pistacia vera) rootstocks to different salt levels after inoculation with arbuscular mycorrhizal fungi Glomus mosseae and compared it with non-mycorrhizal plants (control). Our results confirmed the depressing effect of salt stress on mycorrhization extent and showed that the effect of salinity on colonization rate is completely under the influence of host plant. In this experiment, mycorrhizal symbiosis could enhance plants total dry mass (TDM), electron transfer on the donor and the acceptor side of PSII, decrease the energy dissipation and increase the comprehensive photosynthesis performance under salt stress as well as under normal conditions. We found that both donor and acceptor sides of PSII are the target sides under high salinity in pistachio rootstocks. We also found that performance index is the parameter that better reflects the responses of the studied rootstocks to progressive salt stress. Bane-baqi was less affected by salinity in terms of TDM followed by Sarakhs and Abareqi.     

Growth of mycorrhizal tomato and mineral acquisition under salt stress

 High salt levels in soil and water can limit agricultural production and land development in arid and semiarid regions. Arbuscular mycorrhizal fungi (AMF) have been shown to decrease plant yield losses in saline soils. The objective of this study was to examine the growth and mineral acquisition responses of greenhouse-grown tomato to colonization by the AMF Glomus mosseae [(Nicol. And Gerd.) Gerd. and Trappe] under varied levels of salt. NaCl was added to soil in the irrigation water to give an EC_e of 1.4 (control), 4.7 (medium) and 7.4 dS m^–1 (high salt stress). Plants were grown in a sterilized, low P (silty clay) soil-sand mix. Mycorrhizal colonization was higher in the control than in saline soil conditions. Shoot and root dry matter yields and leaf area were higher in mycorrhizal than in nonmycorrhizal plants. Total accumulation of P, Zn, Cu, and Fe was higher in mycorrhizal than in nonmycorrhizal plants under both control and medium salt stress conditions. Shoot Na concentrations were lower in mycorrhizal than in nonmycorrhizal plants grown under saline soil conditions. The improved growth and nutrient acquisition in tomato demonstrate the potential of AMF colonization for protecting plants against salt stress in arid and semiarid areas. Accepted: 21 February 2000     

低温胁迫下丛枝菌根真菌对玉米光合特性的影响

利用盆栽试验,在15 ℃和5 ℃低温胁迫下研究了丛枝菌根（AM）真菌对玉米生长、叶绿素含量、叶绿素荧光和光合作用的影响.结果表明：低温胁迫抑制了AM真菌的侵染;接种AM真菌的玉米地上部和地下部干物质量、相对叶绿素含量高于不接种植株.与非菌根玉米相比,菌根玉米具有较高的最大荧光（F_m）、可变荧光（F_v）、最大光化学效率（F_v/F_m）和潜在光化学效率（F_v/F_o）及较低的初始荧光（F_o）,并且在5 ℃处理中差异显著.接种AM真菌使玉米叶片的净光合速率（P_n）和蒸腾速率（T_r）显著增强;低温胁迫下,菌根植株的气孔导度（G_s）显著高于非菌根植株;而胞间CO₂浓度（C_i）显著低于非菌根植株.表明AM真菌可通过提高叶绿素含量及改善叶片叶绿素荧光和光合作用来减轻低温胁迫对玉米植株造成的伤害,提高玉米耐受低温的能力,进而提高玉米的生物量,促进玉米生长.     

The effects of salt stress and arbuscular mycorrhiza on plant neighbour effects and self-thinning

Abiotic and biotic factors can alter the nature and strength of plant–plant interactions and therefore self-thinning (density-dependent mortality), but few studies have looked at how such factors interact. We investigated how salt stress and arbuscular mycorrhizal fungi (AMF) influence plant neighbour effects and self-thinning in experimental populations of Medicago sativa. We obtained two mycorrhizal levels by applying the fungicide benomyl (low AMF) or not (high AMF) at three salinity levels (0.05%, 0.2% and 0.5%). In experiment 1, we investigated how salinity and AMF interact to influence plant interaction intensity using a neighbour removal treatment. In experiment 2, we investigated how self-thinning dynamics vary under salinity conditions and different AMF levels at two initial plant densities (6000 and 17,500 seeds m^?2). Shoot biomass and plant density were measured 30, 60 and 90 days after sowing. Standardized major axis regression was used to estimate self-thinning parameters. In experiment 1, AMF increased competitive plant neighbour effects when there was no salinity stress, but this enhancement was not significant with increasing salinity. In experiment 2, there were effects of salinity and AMF on the self-thinning trajectory. The slope of the log (mean shoot biomass per unit area) vs. log density relationship was significantly steeper for the high AMF treatment than for the low AMF treatment without salinity, but the effect of AMF level on the self-thinning exponent was not significant under the two higher salinity levels. The effect of AMF treatments on the intercept of the self-thinning line was not significant at 0.2% salinity but was significant at 0.5% salinity, higher elevation for high AMF treatment. In self-thinning populations, AMF decreased the survival rate without salinity, but increased the survival rate at the highest salinity level. Our results support the hypothesis that salinity and AMF interact to influence plant neighbour effects and self-thinning. Under no-salinity conditions, AMF increased competition, steepened the self-thinning line and decreased survival rate, but these effects of AMF were not significant in the presence of salinity.     

Inoculation with arbuscular mycorrhizal fungi alleviates harmful effects of drought stress on damask rose

This study was conducted to examine the role of arbuscular mycorrhiza fungi (AMF) in alleviating the adverse effects of drought stress on damask rose (Rosa damascena Mill.) plants. Four levels of drought stress (100, 75, 50, and 25% FC) were examined on mycorrhizal and non-mycorrhizal plants in pots filled with sterilized soil. Our results showed that increasing drought stress level decreased all growth parameters, nutrient contents, gas exchange parameters, and water relations indicators. Under different levels of drought stress, mycorrhizal colonization significantly increased all studied parameters. P_n, g_s, and E of the mycorrhizal plants was higher than those of non-mycorrhizal plants under different levels of drought stress. The increase in those rates was proportional the level of the mycorrhizal colonization in the roots of these plants. Majority of growth, nutrition, water status and photosynthetic parameters had a great dependency on the mycorrhizal colonization under all levels of drought stress. The results obtained in this study provide a clear evidence that AMF colonization can enhance growth, flower quality and adaptation of rose plants under different drought stress levels, particularly at high level of drought stress via improving their water relations and photosynthetic status. It could be concluded that colonization with AMF could help plants to tolerate the harmful effects caused by drought stress in arid and semi-arid regions.     

接种AMF对菌根植物和非菌根植物竞争的影响

为了研究丛枝菌根真菌(arbuscular mycorrhizal fungus, AMF)对菌根植物与非菌根植物种间竞争的影响,以玉米(菌根植物)和油菜(非菌根植物)作为供试植物,分别进行间作、尼龙网分隔和单作,模拟这两种植物之间不同的竞争状态,接种丛枝菌根真菌Glomus intraradices和Glomus mosseae,比较菌根植物和非菌根植物的生长和磷营养状况,分析AMF侵染对植物种间竞争作用的影响。结果显示,与单作相比,间作模式下玉米的生物量及磷营养状况有所降低,但其菌根依赖性却有所提高。与不接种相比,接种处理显著降低了间作体系油菜根系的磷含量和磷吸收量,但趋于改善菌根植物玉米的磷营养状况。因此,接种AMF可以降低非菌根植物的磷营养状况及生物量,使得菌根植物的相对竞争能力明显提高,说明AMF在维持物种多样性方面有着重要的作用。