菌根植物适应低磷胁迫的分子机制

丛枝菌根 (AM) 真菌能够和绝大多数陆生植物建立共生体系,对于植物适应低磷胁迫具有重要作用。已有很多研究从不同角度揭示了宿主植物和AM真菌协同适应低磷胁迫的生理机制,并已深入到分子和信号水平。本文归纳了近年来相关研究成果,从磷胁迫信号感知、有机酸分泌、磷酸酶与激素合成相关基因、磷酸盐转运蛋白基因、转录因子与小分子物质miRNA等若干方面讨论了菌根共生体系响应和适应磷胁迫的分子机理,重点介绍了1) 环境磷浓度作为营养信号诱发菌根植物的生理响应过程及其在共生体系建立中的关键作用;2) AM真菌调节植物激素平衡进而影响植物生长发育和根系构型的生理机制;3) 丛枝菌根涉及的植物、真菌以及菌根特异诱导植物产生的磷酸盐转运蛋白基因在磷酸盐摄取中的特殊作用及可能调控机制;4) 转录因子作为感知磷胁迫信号和调控转录表达水平的枢纽,在增强植物适应磷胁迫能力方面的重要贡献。这些因素既单独作用又相互关联,共同构成菌根植物适应磷胁迫的分子调控网络。未来需要着重加强菌根共生界面的磷转运机制、菌根植物适应低磷胁迫的转录因子调节,以及各调控因子相互作用研究,从而全面揭示菌根植物适应低磷胁迫的分子调控网络,为发展和应用菌根技术调控植物磷营养奠定理论基础。     

黄瓜生长素受体同源基因CsAFB和CsTIR在拟南芥中的转化与功能表达

植物生长素受体是生长素信号通路中的重要因子.基于前期克隆得到了黄瓜(Cucumis sativus)生长素受体同源基因生长素信号F-box蛋白基因(auxin signaling F box protein,CsAFB)和转运抑制响应基因(transport inhibitor response,CsTIR),为进一步证实和研究这2个基因的功能,本研究利用拟南芥(Arabidopsis thaliana)Col-0野生型和生长素受体编码基因功能缺陷突变体tirl-1为材料,导入黄瓜生长素受体同源基因,获取纯合转基因系.检测发现,拟南芥突变体tirl-1转入CsAFB/TIR基因后根系发育、外源生长素敏感性和细胞伸长反应均恢复至野生型水平.检测发现,野生型拟南芥中过量表达黄瓜CsAFB/TIR,尤其是Cs TIR基因,植株主根伸长明显受抑,侧根数量剧增,子叶下卷,叶柄上翘,真叶叶缘向离轴面弯曲,顶端优势明显.本研究表明,CsAFB/TIR基因功能类似拟南芥TIR1基因,为黄瓜生长素受体同源基因;过量表达该类基因通过增加生长素受体数量、扩大生长素信号的方式参与调控植物生长发育;为进一步在黄瓜中研究生长素受体功能及作用机理提供理论依据.     

同位素示踪研究丛枝菌根真菌吸收不同氮素并向寄主植物输运的机理

丛枝菌根(Arbuscular mycorrhizae,AM)是自然界中普遍存在的与植物共生的真菌[1]。目前人们对AM真菌的基础研究主要是着眼于与植物共生的生化代谢机理和信号机制、遗传学、建立共生关系相关基因的研究[2-8]。     

矿质养分和激素对根毛生长发育的影响及作用机制

【目的】植物矿质养分和水分的吸收利用赖于根系,根系中根毛的生长发育不仅扩大了根系吸收表面积,促进了矿质养分和水分的吸收还有助于植物根的固定以及与土壤微生物的互作。本文从矿质养分角度(氮、 磷、 钾、 钙、 铁)和激素角度(生长素、 乙烯、 茉莉酸、 独脚金内酯、 油菜素内酯)探讨影响根毛生长发育的因子及作用机理。【主要进展】氮对根毛生长发育的影响与茉莉酸和乙烯有关, 磷与生长素、 乙烯、 独脚金内酯互作调控根毛生长发育;生长素和乙烯以交互作用调控根毛生长发育,茉莉酸、 独角金内酯和油菜素甾醇对根毛生长发育的作用是部分依赖生长素或乙烯途径;植物体内生长素和乙烯等激素的平衡对根毛的生长发育起着重要作用。【建议和展望】基于以上分析,从蛋白激酶及其相关调控基因及转录因子等方面可深入探析矿质养分、 植物激素等对根毛和丛枝菌根生长发育的影响。     

一氧化氮与植物激素的相互作用及其关系

NO在植物的生长发育、生理及信号传递过程中有着重要的调节作用。本文通过从植物根系的生长、种子萌发、程序性细胞死亡、光形态的建成、气孔的关闭及抑制其开放、成熟和衰老等方面对一氧化氮（NO）作为植物激素下游的信号分子发挥的生理功能进行了综述,进而对NO与植物激素生长素、赤霉素、细胞分裂素、脱落酸以及乙烯的相互作用加以讨论,来阐明NO与植物激素之间的关系,并对未来的研究方向作出展望,为NO与植物激素关系的研究提供理论参考。     

烟草与丛枝菌根真菌的共生效应研究进展

丛枝菌根(Arbuscular mycorrhiza,AM)真菌是陆地生态系统中广泛存在的一类专性共生土壤微生物,是根系土壤区域中重要的功能菌群之一.AM真菌可侵染植物根系形成丛枝菌根共生体,改变植物根系形态和改善营养状况,从而提高宿主植物的生长发育、产量、质量和抗逆性.目前从烟草根系土壤分离报道的AM真菌已达13属5...     

脱落酸通过影响生长素合成及分布抑制拟南芥主根伸长

脱落酸(ABA)在介导植物生长发育及逆境响应中发挥重要功能,但ABA抑制根伸长的机制尚不清楚。本文以拟南芥为材料,通过研究ABA对拟南芥根伸长的影响以及ABA受体突变体根发育表型的鉴定,探讨ABA抑制植物主根的机制。研究发现:ABA能够抑制主根生长及伸长,并且经典受体PYR1/PRL介导了ABA抑制根伸长的过程;ABA能够改变细胞周期蛋白CycB1;1::GUS表达模式,并影响根中生长素分布和响应。结果表明,ABA可能通过影响生长素在根部的分布和剂量,进而影响根尖分生区细胞分裂,从而抑制根伸长。     

小麦矮秆突变体DC20的转录组分析

为探明小麦矮杆突变体DC20致矮的分子调控机制,以小麦D6-3(WT)及其经高能混合粒子场诱变处理得到的矮秆突变体DC20为试验材料,通过对突变体DC20和WT的转录组分析,挖掘与DC20致矮相关差异表达基因。结果表明,在株高差异起始的孕穗期茎秆中,DC20与WT之间存在2 153个差异表达基因(DEGs),除参与糖代谢、能量代谢、转录调控、转录后修饰和翻译及翻译后修饰途径外,其中有47个差异表达基因显著富集在植物激素GAs的生物合成和IAA的动态平衡调节及信号转导、细胞周期调控以及细胞伸长等相关途径。大部分差异表达基因表现为表达下调,少量抑制因子的表达量上调。内源植物激素检测结果显示,DC20孕穗期茎秆中的IAA、GA₁和GA₃含量均显著低于WT。表明辐射诱变处理产生的变异是通过植物激素GAs与IAA的协同作用,调控细胞周期以及细胞伸长等途径相关基因的下调表达,从而对DC20的株高产生影响,形成矮化表型。本研究结果为更好地运用辐射诱变育种手段进行作物育种以及阐明矮秆突变体形成的分子调控机理研究提供了一定的理论依据。     

AM真菌与紫云英Ri T-DNA转化根双重培养体系的建立

丛枝菌根(Arbuscular mycorrhiza,AM)真菌是一类古老的植物共生真菌,与植物约有4亿年共生史,能与80%以上的陆生维管植物形成互惠共生体—丛枝菌根,可以改善植物对磷素营养的吸收,增强植物的抗逆性、抗病力或耐病力,与根瘤菌共同作用于豆科植物促进豆科植物正常的生长发育,提高产量,在自然生态系统和实践应用中发挥着重要作用。     

植物磷转运子 PHT1 家族研究进展

【目的】磷是植物生长发育所必需的大量营养元素。植物 PHT1 磷转运蛋白家族在植物磷吸收、运转及再利用等过程中发挥了重要作用。迄今已在多种高等植物中相继分离出大量 PHT1 家族基因。本文综述了国内外关于植物 PHT1 家族的主要研究进展,详细阐述了植物 PHT1 家族的表达模式、功能及可能的调控途径。 主要进展植物 PHT1 家族属于 MFS (major facilitator superfamily) 超家族,不同物种 PHT1 家族蛋白的结构非常保守,通常具有 12 个亲脂跨膜结构域,形成“6 螺旋–亲水大环–6 螺旋”式的结构镶嵌于质膜当中。同时,该家族具有 H₂PO₄–/nH+ 共运子、糖转运子和 MFS 通用转运子等特征结构域和一段保守的氨基酸特征序列 GGDYPLSATIMSE。一般情况,植物 PHT1 家族基因吸收转运 1 个无机磷需要 2～4 个质子协同进入质膜,并伴随膜电位的变化。植物 PHT1 家族的磷转运特性差异较大,其动力学参数 Km 值差别较大。高等植物 PHT1 家族成员众多。在拟南芥、水稻、大豆、茄科植物及其他物种中的研究发现,PHT1 家族各成员间的时空表达模式存在差异,多数成员受低磷信号调控且主要在根部表达,少部分成员在除根以外的其他器官中表达,并行使相应的磷转运功能。已有研究表明,植物 PHT1 家族基因的转录水平受到多因素的调控,例如外界环境中的无机磷浓度,转录因子如 MYB 家族、WRKY 家族以及 ZAT6 等基因能与 PHT1 家族基因启动子区的特殊调控元件如 MYCS 元件、P1BS 元件及 W-box 元件等结合,调控基因的转录。此外,部分 PHT1 家族基因的转录水平受丛枝菌根真菌 (arbuscular mycorrhizal fungi,AMF) 的调控。除了转录水平的调控,关于植物 PHT1 家族转录后水平的调控途径同样取得了较大进展。PHF1 基因、含 SPX 结构域的蛋白家族、MicroRNA、蛋白磷酸化与去磷酸化、染色质修饰及其他等一系列调控途径均参与到 PHT1 家族基因的转录后调控及信号转导。植物激素如生长素、乙烯和细胞分裂素等也参与这一调控过程。 建议与展望植物对磷吸收利用的分子调控机理及信号转导途径十分复杂,因此,培育磷高效利用基因型作物任重而道远。关于植物 PHT1 家族基因的研究已从模式植物向作物及其他高等植物中扩展,然而对该家族蛋白的生化及结构生物学等研究还待进一步深入。同时,对于一些基因组较复杂的多倍体物种如甘蓝型油菜、小麦、大麦及棉花等,仍有待开展进一步研究。     

The role of arbuscular mycorrhizal colonization in the carbon and nutrient economy of the tripartite symbiosis with nodulated Phaseolus vulgaris

In the tripartite symbiosis between nodulated legume roots and arbuscular mycorrhizal (AM) fungi, symbiont sink strength may depend upon developmental stage and the nutrient benefits to the host plant. The cost-benefits of the tripartite symbiosis were investigated in terms of C-economy and nutrition. Nodulated Phaseolus vulgaris seedlings, with and without AM, were hydroponically grown under high (2 mM) and low (1 μM) P conditions in an N-free Long Ashton nutrient solution. Plants were sequentially harvested at 17, 24 and 31 days after emergence. At each harvest, measurements for biomass, N₂-fixation, photosynthesis, root respiration, calculated C and nutritional economy were taken. Nodular growth was suppressed by the early development of AM colonization. This coincided with higher photosynthetic and respiratory rates in AM plants. These effects were most pronounced under low P when AM colonization peaked. Once AM levels reached the plateau phase, the efficiency of P nutrition increased. This was followed by improved nodular and host growth and enhanced N₂-fixation. This indicates that the AM was the dominant symbiont for host C in the tripartite symbiosis, due to its rapid development and subsequent role in supplying P more effectively to both host and nodules.     

Arbuscular mycorrhizae in a long-term field trial comparing low-input (organic, biological) and high-input (conventional) farming systems in a crop rotation

 Arbuscular mycorrhizal (AM) root colonization was studied in a long-term field trial in which four farming systems currently in use in Switzerland were continuously applied to a randomized set of plots at a single field site from 1978 till 1993. There were two low-input farming systems (organic and bio-dynamic) and two high-input (conventional) farming systems (according to Swiss guidelines of integrated plant production with and without farmyard manure). The systems had an identical 7-year crop rotation and tillage scheme and differed essentially only in the amount and type of fertilizer supplied and in plant protection management. The percentage of root colonization by AM fungi was determined in field samples 2–3 times over the growing season in crops in the rotation, namely in winter wheat (Triticum aestivum L. cv. Sardona), vetch-rye and grass-clover. We found the percentage of root length colonized by AM fungi to be 30–60% higher (P≤0.05) in the plants grown in soils from the low-input farming systems than in those grown in conventionally farmed soils. Approximately 50% of the variation of AM root colonization was explained by chemical properties of the soils (pH, soluble P and K, exchangeable Mg), the effect of soluble soil P being most pronounced. The potential of the field soils from the differently managed plots to cause symbiosis with AM fungi was tested in a glasshouse experiment, using wheat as a host plant. Soils from the low-input farming systems had a greatly enhanced capacity to initiate AM symbiosis. The relative differences in this capacity remained similar when propagules of the AM fungus Glomus mosseae were experimentally added to the soils, although overall root colonization by AM fungi was 2.8 times higher. Received: 27 August 1999     

免耕和秸秆覆盖对黑垆土磷素形态组分的影响

[目的]探究免耕及添加秸秆条件下黑垆土土壤磷组分特征及其与AM真菌侵染的关系,了解雨养农业区农业系统磷素利用效率。[方法]在陇东黄土高原黑垆土区域,测定传统耕作、传统耕作+秸秆覆盖、免耕和免耕+秸秆覆盖4种处理小麦—玉米—大豆轮作系统中玉米阶段土壤全磷、速效磷组分及AM真菌菌根侵染率。[结果]水土保持耕作处理实施9a后,免耕和秸秆覆盖处理下0—5cm土壤磷素含量显著提高,活性磷组分H2O—Pi,NaHCO3—Pi,NaOH—Pi分别比对照提高84.6%,85.2%和56.6%;活性无机磷(H2O—Pi,NaHCO3—Pi之和)和潜在活性磷(NaOH—Pi)分别占总无机磷的11.4%和4.5%,全磷含量与磷组分、速效磷与磷组分呈显著正相关,2个免耕处理菌根侵染率分别比对照增加20.8%和16.5%。[结论]免耕和秸秆覆盖显著提高了土壤磷含量,免耕对AM真菌菌根侵染率有积极影响。     

Arbuscular mycorrhizal fungi regulate plant mineral nutrient uptake and partitioning in iron ore tailings undergoing eco-engineered pedogenesis

Excess available K and Fe in Fe ore tailings with organic matter amendment and water-deficiencies may restrain plant colonization and growth, which hinders the formation of eco-engineered soil from these tailings for sustainable and cost-effective mine site rehabilitation. Arbuscular mycorrhizal (AM) fungi are widely demonstrated to assist plant growth under various unfavorable environments. However, it is still unclear whether AM symbiosis in tailings amended with different types of plant biomass and under different water conditions could overcome the surplus K and Fe stress for plants in Fe ore tailings, and if so, by what mechanisms. Here, host plants (Sorghum sp. Hybrid cv. Silk), either colonized or noncolonized by the AM fungi (Glomus spp.), were cultivated in lucerne hay (LH, C:N ratio of 18)- or sugarcane mulch (SM, C:N ratio of 78)-amended Fe ore tailings under well-watered (55% water-holding capacity (WHC) of tailings) or water-deficient (30% WHC of tailings) conditions. Root mycorrhizal colonization, plant growth, and mineral elemental uptake and partitioning were examined. Results indicated that AM fungal colonization improved plant growth in tailings amended with plant biomass under water-deficient conditions. Arbuscular mycorrhizal fungal colonization enhanced plant mineral element uptake, especially P, both in the LH- and SM-amended tailings regardless of water condition. Additionally, AM symbiosis development restrained the translocation of excess elements (i.e., K and Fe) from plant roots to shoots, thereby relieving their phytotoxicity. The AM fungal roles in P uptake and excess elemental partitioning were greater in LH-amended tailings than in SM-amended tailings. Water deficiency weakened AM fungal colonization and functions in terms of mineral element uptake and partitioning. These findings highlighted the vital role AM fungi played in regulating plant growth and nutrition status in Fe ore tailings technosol, providing an important basis for involvement of AM fungi in the eco-engineered pedogenesis of Fe ore tailings.     

Effects of soil application of olive mill wastewaters on the structure and function of the community of arbuscular mycorrhizal fungi

Recycling of olive mill wastewaters (OMW) into agricultural soils is a controversial issue since benefits to soil fertility should counterbalance potential short-term toxicity effects. We investigated the short-term effects of OMW on the soil-plant system, regarding the diversity, structure and root colonization capacity of arbuscular mycorrhizal (AM) fungi and the respective growth response of Vicia faba L, commonly used as green manure in olive-tree plantations. A compartmentalized pot system was used that allowed the establishment of an AM fungal community in one compartment (feeder) and the application of three OMW dose levels in an adjacent second compartment (receiver). At 0, 10, and 30 days after OMW treatment (DAT), V. faba pre-germinated seeds were seeded in the receiver compartment. At harvest, shoot and root dry weights, AM fungal root colonization, soil hyphal length and P availability were recorded in the receiver compartment. In addition, OMW effects on AM fungal diversity in plant roots were studied by DGGE. A transient effect of OMW application was observed; plant growth and AM fungal colonization were initially inhibited, whereas soil hyphal length was stimulated, but in most cases differences were absent when seeding was performed 30 DAT. Similarly, changes induced in the structure of the root AM fungal community were of transient nature. Cloning and sequencing of all the major DGGE bands showed that roots were colonized by Glomus spp. The transient effects of OMW on the structure and function of AM fungi could be attributed to OMW-derived phytoxicity to V. faba plants or to an indirect effect via alteration of soil nutritional status. The high OMW dose significantly increased soil P availability in the presence of AM fungi, suggesting efficient involvement of AM fungi in organic-P minerilization. Overall our results indicate that soil application of OMW would cause transient changes in the AM fungal colonization of V. faba plants, which, would not impair their long-term plant growth promoting ability.     

Arbuscular Mycorrhizal Mediated Nutrition in Plants

ABSTRACT

Arbuscular mycorrhizae (AM) are the symbiotic fungi that predominate in the roots and soils of agricultural crop plants. The most recognized beneficial effect of these fungi is to enhance host plant uptake of relatively immobile nutrients, in particular phosphorus (P), and several micronutrients. The AM fungi absorb inorganic P either from the soluble P pools in the soil, or from insoluble forms such as rock phosphates as well as from insoluble organic sources. Recent studies show that mycorrhizal fungi would have access to rock phosphate through localized alterations of pH and/or by the production of organic acid anions that may act as chelating agents. The AM colonization also improves plant N nutrition. Generally mycorrhizal symbiosis more influences on nitrogen (N) uptake and translocation if ammonium (NH₄ ⁺) rather than nitrate (NO₃ ^?) is the nitrogen source. However, under drought stress the role of mycorrhizae in NO₃ ^? transport to the root surface may be significant as the NO₃ ^? mobility is severely restricted due to its low concentration and diffusion rate under such circumstances. However, as yet little is known about the mechanism of N uptake by the AM fungi. Uptake of micronutrients is also influenced by mycorrhizal colonization.     

Impact of soil nitrogen concentration on Glomus spp.-Sinorhizobium interactions as affecting growth, nitrate reductase activity and protein content of Medicago sativa

Our objective was to evaluate how increasing levels of N in the medium (0, 4, 8 and 16 mmol N added kg^-1 soil) affect the interaction between Sinorhizobium and arbuscular mycorrhiza (AM) fungi in the tripartite symbiosis with Medicago sativa. Growth response, nutrient acquisition, protein content, and nitrate reductase (NR) activity were measured both in plant shoots and roots. Results showed that N levels in soil did not affect mycorrhizal colonization but they strongly influenced nodulation, particularly of mycorrhizal plants. Mycorrhizal colonization was required for a proper nodulation when no N was applied to soil. In contrast, the addition of 4 mmol N kg^-1 soil reduced nodulation only in mycorrhizal plants and 8 mmol N added kg^-1 soil allowed nodule formation only in non-mycorrhizal plants. Nodulation was totally inhibited in all treatments with the addition of 16 mmol N added kg^-1 soil. N addition enhanced NR activity in all the treatments, while AM colonization increased the proportion of NR allocated to roots. This effect was more pronounced under the lowest N levels in the medium. The two AM fungal species showed different distribution pattern of enzymatic activities in plant tissues indicating specific physiological traits. Protein content as well as the relative proportion of protein in roots were greatly increased after mycorrhizal colonization. Glomus intraradices-colonized plants had the highest protein content in shoot and root. Mycorrhizal effects on growth, N acquisition and biochemical variables cannot be interpreted as an indirect P-mediated effect since P content was lower in mycorrhizal plants than in those which were P fertilized. Mycorrhizal colonization increased the N content in plants irrespective of the N level, but the effectiveness of AM fungi on plant N acquisition depended on the AM fungus involved, G. intraradices being the most effective, particularly at the highest N rate. N₂ fixation, enhanced by AM colonization, contributed to N acquisition when a moderate N quantity was available in the soil. Nevertheless, under a high N amount the nodulating process and/or fixing capacity by Sinorhizobium was reduced in AM plants. In contrast, the AM fungal mycelium from a particular mycorrhizal fungus may continue to contribute efficiently to the N uptake from the soil even at high N levels. These results demonstrate the particular sensitivity of AM fungal species in terms of their growth and/or function to increasing N amounts in the medium. A selection of AM fungi used to address specific environmental conditions, such as N fertilization regimes comparable to those used in agronomic practices, is required for a better use of N applied to soil.     

Effects of salinity and flooding on the infectivity of salt marsh arbuscular mycorrhizal fungi in<Emphasis Type="Italic"> Aster tripolium</Emphasis> L.

Salt marshes are characterized by the occurrence of combined salinity and flooding stresses. The individual and combined effects of salinity and flooding on the establishment and activity of arbuscular mycorrhizal (AM) colonization in the salt marsh halophyte Aster tripolium L. by indigenous salt marsh AM fungi were evaluated. A. tripolium plants were cultivated in a mixture of sand and salt marsh soil under different salinity concentrations (5%, 50% or 100% artificial seawater) and water regimes (non-flooding, tidal flooding and continuous flooding). Plants were harvested after 3 and 8 weeks and their growth was negatively influenced by increased salinity and water level. Increased salinity level affected the establishment of AM colonization, AM fungal growth and activity (measured as succinate dehydrogenase activity) within roots, and extraradical mycelium growth. The influence of flooding on the establishment of colonization and on intra- and extraradical AM fungal growth was dependent on the water regime. Continuous flooding reduced colonization and AM fungal growth, whereas tidal flooding did not affect these parameters unless combined with intermediate salinity level (50% seawater) at the end of the experiment. The water regime did not influence AM active colonization. The ratio of root to soil AM fungal growth increased as the water level increased. The results of this study demonstrate that the establishment and activity of AM colonization in A. tripolium is more influenced by salinity than by flooding, and suggests that the functionality of salt marsh AM fungi is not affected by flooding.     

铬稳定同位素在丛枝菌根蒲公英中的分馏及根外菌丝对铬的吸收

As common soil fungi that form symbioses with most terrestrial plants,arbuscular mycorrhizal(AM) fungi play an important role in plant adaptation to chromium(Cr) contamination.However,little information is available on the underlying mechanisms of AM symbiosis on plant Cr resistance.In this study,dandelion(Taraxacum platypecidum Diels.) was grown with and without inoculation of the AM fungus Rhizophagus irregularis and Cr uptake by extraradical mycelium(ERM) was investigated by a compartmented cultivation system using a Cr stable isotope tracer.The results indicated that AM symbiosis increased plant dry weights and P concentrations but decreased shoot Cr concentrations.Using the Cr stable isotope tracer technology,the work provided possible evidences of Cr uptake and transport by ERM,and confirmed the enhancement of root Cr stabilization by AM symbiosis.This study also indicated an enrichment of lighter Cr isotopes in shoots during Cr translocation from roots to shoots in mycorrhizal plants.