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.     

Regulation of the Nitrogen Transfer Pathway in the Arbuscular Mycorrhizal Symbiosis: Gene Characterization and the Coordination of Expression with Nitrogen Flux

The arbuscular mycorrhiza (AM) brings together the roots of over 80% of land plant species and fungi of the phylum Glomeromycota and greatly benefits plants through improved uptake of mineral nutrients. AM fungi can take up both nitrate and ammonium from the soil and transfer nitrogen (N) to host roots in nutritionally substantial quantities. The current model of N handling in the AM symbiosis includes the synthesis of arginine in the extraradical mycelium and the transfer of arginine to the intraradical mycelium, where it is broken down to release N for transfer to the host plant. To understand the mechanisms and regulation of N transfer from the fungus to the plant, 11 fungal genes putatively involved in the pathway were identified from Glomus intraradices, and for six of them the full-length coding sequence was functionally characterized by yeast complementation. Two glutamine synthetase isoforms were found to have different substrate affinities and expression patterns, suggesting different roles in N assimilation. The spatial and temporal expression of plant and fungal N metabolism genes were followed after nitrate was added to the extraradical mycelium under N-limited growth conditions using hairy root cultures. In parallel experiments with ¹⁵N, the levels and labeling of free amino acids were measured to follow transport and metabolism. The gene expression pattern and profiling of metabolites involved in the N pathway support the idea that the rapid uptake, translocation, and transfer of N by the fungus successively trigger metabolic gene expression responses in the extraradical mycelium, intraradical mycelium, and host plant.The arbuscular mycorrhizal (AM) symbiosis brings together the roots of the majority of land plant species and fungi of the phylum Glomeromycota to great mutual advantage (Smith and Read, 2008). AM fungi improve the acquisition of phosphate, nitrogen (N), sulfur, and trace elements such as copper and zinc (Clark and Zeto, 2000; Allen and Shachar-Hill, 2008) and increase the biotic and abiotic stress resistance of their host (Smith et al., 2010). In return, the host transfers up to 20% of its photosynthetically fixed carbon to the AM fungus (Jakobsen and Rosendahl, 1990), which depends on its host plant for its carbon supply (Bago et al., 2000).N is the nutrient whose availability most commonly limits plant growth in natural ecosystems. AM fungi can take up NO₃⁻NH₄⁺ and can also increase access to organic N sources from the soil (Ames et al., 1983; Johansen et al., 1993; Bago et al., 1996; Hodge et al., 2001). The translocation by the fungus can represent a significant route for N uptake by the plant (Johansen and Jensen, 1996). For example, Toussaint et al. (2004) showed that in an in vitro mycorrhiza at least 21% of the total N uptake in the roots came from the fungal extraradical mycelium (ERM); for other mycorrhizal systems, even larger proportions have been described (more than 30% and 50%; Govindarajulu et al., 2005; Jin et al., 2005). Tanaka and Yano (2005) reported that 75% of the N in leaves of mycorrhizal maize (Zea mays) was taken up by the ERM of Glomus aggregatum.A mechanism of N transfer from the fungus to the plant has been proposed (Bago et al., 2001) that involves the operation of a novel metabolic route in which N was translocated from the ERM to the intraradical mycelium (IRM) as Arg but transferred to the plant without carbon as inorganic N. This mechanism has been supported by labeling experiments (Johansen et al., 1996; Govindarajulu et al., 2005; Jin et al., 2005), enzyme activity analysis (Cruz et al., 2007), and limited gene expression data (Govindarajulu et al., 2005; Gomez et al., 2009; Guether et al., 2009). Nevertheless, our molecular knowledge of the metabolic and transport pathways involved and how they are regulated is still rudimentary. A better understanding of the mechanism and regulation of N uptake assimilation, translocation, and transfer to the host is important for potential applications of AM fungi as biofertilizers, bioprotectors, and bioregulators in sustainable agriculture and restoration as well as for understanding the role of AM fungi in natural ecosystems (Bruns et al., 2008).In this study, we postulate that the uptake, translocation, and transfer of N by the fungus triggers the metabolic gene expression responses successively in the ERM, IRM, and host plant, which will result in the synthesis and accumulation of Arg in the ERM, the turnover of Arg to release ammonium in the IRM, and the assimilation of ammonium by the host plant via the glutamine synthetase (GS)/glutamate synthase (GOGAT) pathway inside the root (Fig. 1). To test these predictions, 11 genes involved in the N primary assimilation and metabolism were cloned and verified from Glomus intraradices; six enzymes with full-length coding sequences (CDSs) were functionally characterized by yeast knockout mutant complementation. Two GS proteins were found to have different substrate affinities and expression patterns, suggesting that they have different roles in N assimilation. The time courses of gene expression and N movement in fungal and host tissues were analyzed following nitrate supply to the ERM of a mycorrhiza grown under N-limited conditions. The results substantially increase our knowledge of the identity and regulation of most of the metabolic and transport genes involved in N movement through the AM symbiosis.Open in a separate windowFigure 1.Working model of N transport and metabolism in the symbiosis between plant roots and arbuscular mycorrhizal fungi. N moves (black arrows) from the soil into the fungal ERM, through a series of metabolic conversion reactions into Arg, which is transported into the IRM within the root (host) and there is broken down; N is transferred to and assimilated by the host as ammonia. Red circles refer to the sites of action of the genes identified and analyzed in this study. Blue arrows indicate mechanisms hypothesized to regulate gene expression by N metabolites involved in the pathway.     

接种丛枝菌根（AM）真菌对植物DBP污染的影响

在温室进行盆栽试验,以DBP(邻苯二甲酸二丁酯)为研究对象,以豇豆(Pigna sinensis)为宿主植物,分别接种AM真菌Acaulospora lavis(光壁无梗球囊霉)和Glomus caledonium(苏格兰球囊霉),观察接种AM真菌对植物DBP污染变化的影响．结果表明,接种AM真菌明显控制了植物对DBP的吸收,降低了植物体内DBP的浓度．在低浓度DBP(4mg.kg^-1)土壤处理时,接种Acaulospora lavis和Glomus caledo-nium分别使植物体内DBP浓度比不接种(CK)最大下降32．7％和21．7％;高浓度DBP(100mg.kg^-1)土壤处理时,分别比CK最大下降30．5％和30．0％．接种AM真菌还抑制了DBP由植物根系向地上部的迁移,对减轻植物遭受DBP污染起了一定的作用．     

Arbuscular mycorrhizal fungi can transfer substantial amounts of nitrogen to their host plant from organic material

Nitrogen (N) capture by arbuscular mycorrhizal (AM) fungi from organic material is a recently discovered phenomenon. This study investigated the ability of two Glomus species to transfer N from organic material to host plants and examined whether the ability to capture N is related to fungal hyphal growth. Experimental microcosms had two compartments; these contained either a single plant of Plantago lanceolata inoculated with Glomus hoi or Glomus intraradices, or a patch of dried shoot material labelled with (15)N and (13)carbon (C). In one treatment, hyphae, but not roots, were allowed access to the patch; in the other treatment, access by both hyphae and roots was prevented. When allowed, fungi proliferated in the patch and captured N but not C, although G. intraradices transferred more N than G. hoi to the plant. Plants colonized with G. intraradices had a higher concentration of N than controls. Up to one-third of the patch N was captured by the AM fungi and transferred to the plant, while c. 20% of plant N may have been patch derived. These findings indicate that uptake from organic N could be important in AM symbiosis for both plant and fungal partners and that some AM fungi may acquire inorganic N from organic sources.     

丛枝菌根在植物修复重金属污染土壤中的作用

丛枝菌根（Arbuscular mycorrhizae，AM）是自然界中分布最广的一类菌根，AM真菌能与陆地上绝大多数的高等植物共生，常见于包括重金属污染土壤在内的各种生境中。在重金属污染条件下，AM真菌可以减轻重金属对植物的毒害，影响植物对重金属的吸收和转运，在重金属污染土壤的植物修复中显示出极大的应用潜力。重点介绍了AM真菌对植物重金属耐性的影响及其在植物提取和植物稳定中的应用等方面的进展，讨论了未来研究所面临的任务和挑战。     

Mycorrhizal mediation of plant N acquisition and residue decomposition: Impact of mineral N inputs

Mycorrhizas are ubiquitous plant–fungus mutualists in terrestrial ecosystems and play important roles in plant resource capture and nutrient cycling. Sporadic evidence suggests that anthropogenic nitrogen (N) input may impact the development and the functioning of arbuscular mycorrhizal (AM) fungi, potentially altering host plant growth and soil carbon (C) dynamics. In this study, we examined how mineral N inputs affected mycorrhizal mediation of plant N acquisition and residue decomposition in a microcosm system. Each microcosm unit was separated into HOST and TEST compartments by a replaceable mesh screen that either prevented or allowed AM fungal hyphae but not plant roots to grow into the TEST compartments. Wild oat (Avena fatua L.) was planted in the HOST compartments that had been inoculated with either a single species of AM fungus, Glomus etunicatum, or a mixture of AM fungi including G. etunicatum. Mycorrhizal contributions to plant N acquisition and residue decomposition were directly assessed by introducing a mineral ¹⁵N tracer and ¹³C‐rich residues of a C₄ plant to the TEST compartments. Results from ¹⁵N tracer measurements showed that AM fungal hyphae directly transported N from the TEST soil to the host plant. Compared with the control with no penetration of AM fungal hyphae, AM hyphal penetration led to a 125% increase in biomass ¹⁵N of host plants and a 20% reduction in extractable inorganic N in the TEST soil. Mineral N inputs to the HOST compartments (equivalent to 5.0 g N m^?2 yr^?1) increased oat biomass and total root length colonized by mycorrhizal fungi by 189% and 285%, respectively, as compared with the no‐N control. Mineral N inputs to the HOST plants also reduced extractable inorganic N and particulate residue C proportion by 58% and 12%, respectively, in the corresponding TEST soils as compared to the no‐N control, by stimulating AM fungal growth and activities. The species mixture of mycorrhizal fungi was more effective in facilitating N transport and residue decomposition than the single AM species. These findings indicate that low‐level mineral N inputs may significantly enhance nutrient cycling and plant resource capture in terrestrial ecosystems via stimulation of root growth, mycorrhizal functioning, and residue decomposition. The long‐term effects of these observed alterations on soil C dynamics remain to be investigated.     

Transport of nitrogen and zinc to rhodes grass by arbuscular mycorrhiza and roots as affected by different nitrogen sources (NH<Subscript>4</Subscript><Superscript>+</Superscript>-N and NO<Subscript>3</Subscript><Superscript>−</Superscript>-N)

We investigated the effect of mineral nitrogen forms on transfer of nitrogen (N) and zinc (Zn) from attached compartments to rhodes grass (Chloris gayana) colonised with arbuscular mycorrhizal fungi (AMF). After being pre-cultivated in substrates with adequate nutrient supply and either AMF inoculated (+AM) or left non-inoculated (?AM), rhodes grass was positioned adjacent to an outer compartment holding a similar substrate but applied with labelled nitrogen (¹⁵N) either as ammonium (NH₄ ⁺) or nitrate (NO₃ ^?), and a high supply of Zn (150 mg kg^?1 DS). Plant roots together with fungal mycelium were either allowed to explore the outer compartment (with root access) or only mycorrhizal hyphae were allowed (without root access). Within each access treatment, biomasses of rhodes grass were not significantly affected by AMF inoculation or N form. AMF contribution to plant ¹⁵N uptake was about double in NH₄ ⁺ compared with NO₃ ^?-supplied treatments while the mycorrhizal influence on plant Zn uptake was insignificant. Without root access, the shoot ¹⁵N/Zn concentration ratio was up to ten-fold higher in +AM than –AM treatments and this ratio increase was clearly more pronounced in NH₄ ⁺ than NO₃ ^?-supplied treatments. In conclusion, rhodes grass in symbiosis with the tested AMF acquired more N when supplied with ammonium. Moreover, there is clear indication that although the AMF have transported both nutrients (N and Zn), N was preferentially transferred as compared to Zn. We confirmed that, while rhodes grass is not able to prevent excessive Zn uptake via roots under conditions of high Zn, mycorrhiza is able to avoid excessive Zn supply to the host plant when the fungus alone has access to contaminated patches.     

Phosphate uptake, transport and transfer by the arbuscular mycorrhizal fungus Glomus intraradices is stimulated by increased carbohydrate availability

* The influence of carbohydrate availability to mycorrhizal roots on uptake, metabolism and translocation of phosphate (P) by the fungus was examined in axenic cultures of transformed carrot (Daucus carota) roots in symbiosis with Glomus intraradices. * 14C-labelled carbohydrates, 33P-phosphate and energy dispersive X-ray microanalysis were used to follow the uptake and transfer of C and P in the arbuscular mycorrhizal (AM) symbiosis. * The uptake of P by the extraradical mycelium (ERM) and its translocation to the mycorrhizal roots was stimulated and the metabolic and spatial distribution of P within the fungus were altered in response to increased carbohydrate availability. Sucrose supply resulted in a decrease of polyphosphates and an increased incorporation into phospholipids and other growth-related P pools and also caused elevated cytoplasmic P levels in the intraradical mycelium (IRM) within the root and higher cytoplasmic P levels in the root cortex. * These findings indicate that the uptake of P by the fungus and its transfer to the host is also stimulated by the transfer of carbon from plant to fungus across the mycorrhizal interface.     

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.
     

Transport of radiocaesium by arbuscular mycorrhizal fungi to Medicago truncatula under in vitro conditions

The capacity of arbuscular mycorrhizal (AM) fungi to take up and translocate radiocaesium (Cs) to their host has been shown using the root-organ culture (ROC) system. However, the absence of photosynthetic tissues, lack of a normal root hormonal balance and incomplete source-sink relationships may bias the bidirectional transfer of elements at the symbiotic interface and complicate transport studies. Accordingly, we developed a novel culture system [i.e. the Arbuscular Mycorrhizal-Plant (AM-P) in vitro culture system], where AM fungi and an autotrophic host plant develop under strict in vitro conditions. With this system, we unambiguously demonstrated the capacity of AM fungi to transport Cs. The extraradical fungal hyphae took up 21.0% of the initial supply of 134Cs. Translocation to the plant represented 83.6% of the 134Cs taken up. Distribution of 134Cs in the host plant was 89.8% in the mycorrhizal roots and 10.2% in the shoot. These results confirm that AM fungi can take up, translocate and accumulate Cs. They further demonstrate unambiguously and for the first time that Cs can be transferred from AM fungi to host tissues. These results suggest a potential involvement of AM fungi in Cs biogeochemical cycle and in plant Cs accumulation.     

Nutrient uptake in mycorrhizal symbiosis

The role of mycorrhizal fungi in acquisition of mineral nutrients by host plants is examined for three groups of mycorrhizas. These are; the ectomycorrhizas (ECM), the ericoid mycorrhizas (EM), and the vesicular-arbuscular mycorrhizas (VAM). Mycorrhizal infection may affect the mineral nutrition of the host plant directly by enhancing plant growth through nutrient acquisition by the fungus, or indirectly by modifying transpiration rates and the composition of rhizosphere microflora. A capacity for the external hyphae to take up and deliver nutrients to the plant has been demonstrated for the following nutrients and mycorrhizas; P (VAM, EM, ECM), NH₄ ⁺ (VAM, EM, ECM), NO₃ ^- (ECM), K (VAM, ECM), Ca (VAM, EM), SO₄ ^2- (VAM), Cu (VAM), Zn (VAM) and Fe (EM). In experimental chambers, the external hyphae of VAM can deliver up to 80% of plant P, 25% of plant N, 10% of plant K, 25% of plant Zn and 60% of plant Cu. Knowledge of the role of mycorrhiza in the uptake of nutrients other than P and N is limited because definitive studies are few, especially for the ECM. Although further quantification is required, it is feasible that the external hyphae may provide a significant delivery system for N, K, Cu and Zn in addition to P in many soils. Proposals that ECM and VAM fungi contribute substantially to the Mg, B and Fe nutrition of the host plant have not been substantiated. ECM and EM fungi produce ectoenzymes which provide host plants with the potential to access organic N and P forms that are normally unavailable to VAM fungi or to non mycorrhizal roots. The relative contribution of these nutrient sources requires quantification in the field. Further basic research, including the quantification of nutrient uptake and transport by fungal hyphae in soil and regulation at the fungal-plant interface, is essential to support the selection and utilization of mycorrhizal fungi on a commercial scale.     

Trichoderma harzianum might impact phosphorus transport by arbuscular mycorrhizal fungi

Trichoderma sp. is a biocontrol agent active against plant pathogens via mechanisms such as mycoparasitism. Recently, it was demonstrated that Trichoderma harzianum was able to parasitize the mycelium of an arbuscular mycorrhizal (AM) fungus, thus affecting its viability. Here, we question whether this mycoparasitism may reduce the capacity of Glomus sp. to transport phosphorus ((33)P) to its host plant in an in vitro culture system. (33)P was measured in the plant and in the fungal mycelium in the presence/absence of T. harzianum. The viability and metabolic activity of the extraradical mycelium was measured via succinate dehydrogenase and alkaline phosphatase staining. Our study demonstrated an increased uptake of (33)P by the AM fungus in the presence of T. harzianum, possibly related to a stress reaction caused by mycoparasitism. In addition, the disruption of AM extraradical hyphae in the presence of T. harzianum affected the (33)P translocation within the AM fungal mycelium and consequently the transfer of (33)P to the host plant. The effects of T. harzianum on Glomus sp. may thus impact the growth and function of AM fungi and also indirectly plant performance by influencing the source-sink relationship between the two partners of the symbiosis.     

P metabolism and transport in AM fungi

The arbuscular mycorrhizal symbiosis is mutualistic, based on reciprocal transfer of P from the fungus to the plant and carbon from the plant to the fungus. Thus P is a most important `currency' in the symbiosis. After absorbing P from the soil solution, the fungi first incorporate it into the cytosolic pool, and the excess P is transferred to the vacuoles. The vacuolar P pool probably plays a central role in P supply to the plant. The main forms of inorganic P in fungal vacuoles are orthophosphate and polyphosphate, but organic P molecules may also be present. Long distance translocation of P from the site of uptake in the external mycelium to the site of transfer to the plant is probably achieved via transfer of vacuolar components. This transport would be mediated either by protoplasmic streaming or the motile tubular vacuole-like system. The site of release of P into the interfacial apoplast and thence to the plant is most probably the fungal arbuscules. The biochemical and biophysical processes involved in P metabolism and transfer between cellular compartments in the symbiosis are currently not well understood. Some recent investigations of substrate specificities of phosphatase-type enzymes in AM fungi and other eukaryotic microorganisms, however, have shed new light on earlier results and permit the construction of a hypothetical scheme of P-flow, including possible regulatory factors. Steps in this scheme are experimentally testable and should stimulate future research.     

Phosphorus uptake by a community of arbuscular mycorrhizal fungi in jarrah forest

Communities of indigenous arbusuclar mycorrhizal (AM) fungi are expected to alter phosphorus uptake and biomass productivity of plants according to characteristics of the life cycles of the fungi present and the way they interact with each other inside roots and with host plants. Differences in the relative abundance of AM fungi inside roots could influence P uptake if the fungi present differ in effectiveness at accessing P and transferring it to the plant. However, it is difficult to assess the contribution of AM fungi under field conditions. We investigated P uptake, from point sources of P placed 2, 4 and 6 cm from roots, by plants colonised by a community of AM fungi in jarrah forest soil. Roots were retained within a mesh bag to prevent them from growing towards the point source of P. The relative abundance of morphotypes of fungi inside roots and the P status of plants were assessed after 12 and 16 weeks. First, a bioassay was carried out in undisturbed forest soil cores using two host plants, a forest understorey plant Phyllanthus calycinus Labill and the annual pasture species subterranean clover (Trifolium subterraneaumL.), to assess the infectivity of the indigenous community of AM fungi. Roots of both bioassay host plants were colonised in similar proportions by morphotypes of AM fungi resembling Glomus, Acaulospora, Scutellospora and fine endophytes. In this bioassay, there were positive correlations between the proportion of root length colonised and plant biomass and P uptake for P. calycinus, but not for subterranean clover. In the experiment assessing the capacity of P. calycinus to access P placed at increasing distances from the root, shoot P content and concentration in P. calycinus were greater when P was placed 2 cm compared with 4 and 6 cm from roots. The length of hyphae in the vicinity of the point source of P decreased with increasing distance from the plant. The extent to which the individual AM fungi were involved in P uptake is not known. The Glomus morphotype was dominant at both times of sampling.     

Growth, respiration and nutrient acquisition by the arbuscular mycorrhizal fungus Glomus mosseae and its host plant Plantago lanceolata in cooled soil

Although plant phosphate uptake is reduced by low soil temperature, arbuscular mycorrhizal (AM) fungi are responsible for P uptake in many plants. We investigated growth and carbon allocation of the AM fungus Glomus mosseae and a host plant (Plantago lanceolata) under reduced soil temperature. Plants were grown in compartmented microcosm units to determine the impact on both fungus and roots of a constant 2.7 °C reduction in soil temperature for 16 d. C allocation was measured using two (13)CO(2) pulse labels. Although root growth was reduced by cooling, AM colonization, growth and respiration of the extraradical mycelium (ERM) and allocation of assimilated (13)C to the ERM were all unaffected; the frequency of arbuscules increased. In contrast, root respiration and (13)C content and plant P and Zn content were all reduced by cooling. Cooling had less effect on N and K, and none on Ca and Mg content. The AM fungus G. mosseae was more able to sustain activity in cooled soil than were the roots of P. lanceolata, and so enhanced plant P content under a realistic degree of soil cooling that reduced plant growth. AM fungi may therefore be an effective means to promote plant nutrition under low soil temperatures.     

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.     

植物菌根共生磷酸盐转运蛋白

大多数植物能和丛枝菌根（arbuscular mycorrhiza, AM）真菌形成菌根共生体。AM能够促进植物对土壤中矿质营养的吸收,尤其是磷的吸收。磷的吸收和转运由磷酸盐转运蛋白介导。总结了植物AM磷酸盐转运蛋白及其结构特征,分析其分类及系统进化,并综述了AM磷酸盐转运蛋白介导的磷的吸收和转运过程及其基因的表达调控。植物AM磷酸盐转运蛋白属于Pht1家族成员,它不仅对磷的吸收和转运是必需的,而且对AM共生也至关重要,为进一步了解菌根形成的分子机理及信号转导途径提供了理论基础。     

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。     

Transport of Ferrocyanide by Two Eucalypt Species and Sorghum

The wastes from some industrial processes and the tailings from gold mining contain elevated concentrations of cyanide, which reacts with iron in the media to form iron cyanide complexes. This research examined the transport and possible metabolism of ferrocyanide by two native Australian trees, blue mallee and sugar gum, and by sorghum. Hydroponic studies using ¹⁵N-labeled ferrocyanide showed that both tree species transported ferrocyanide into roots and displayed significant increases in ¹⁵N enrichment and concentration with no evidence of phytotoxicity. A subsequent experiment with blue mallee and membrane-transport inhibitors showed that ¹⁵N enrichment was significantly inhibited in the presence of the protonophore carbonyl cyanide m-chlorophenylhydrazone, suggesting that ferrocyanide uptake is mediated partly by H⁺-symporters. A study of the time dependence of ¹⁵N translocation showed a rapid equilibration of ¹⁵N from ferrocyanide in the root of blue mallee, accompanied by a slow increase in shoot ¹⁵N, suggestive of the metabolism of ferrocyanide in plant roots. A similar experiment with sorghum showed a more rapid translocation of ¹⁵N, suggesting that the transport and/or metabolism of ferrocyanide by roots of this species may differ. The results offer additional incentive for the use of these species as vegetative cover over cyanidation wastes and for cyanide phytoremediation.     

Gibberellin regulates infection and colonization of host roots by arbuscular mycorrhizal fungi

Arbuscular mycorrhiza (AM) is established by the entry of AM fungi into the host plant roots and the formation of symbiotic structures called arbuscules. The host plant supplies photosynthetic products to the AM fungi, which in return provide phosphate and other minerals to the host through the arbuscules. Both partners gain great advantages from this symbiotic interaction, and both regulate AM development. Our recent work revealed that gibberellic acids (GAs) are required for AM development in the legume Lotus japonicus. GA signaling interact with symbiosis signaling pathways, directing AM fungal colonization in host roots. Expression analysis showed that genes for GA biosynthesis and metabolism were induced in host roots around AM fungal hyphae, suggesting that the GA signaling changes with both location and time during AM development. The fluctuating GA concentrations sometimes positively and sometimes negatively affect the expression of AM-induced genes that regulate AM fungal infection and colonization.