联合固氮的研究进展

1975年,Dbereiner实验发现与禾本科植物联合共生的固氮菌并提出根际联合固氮的概念。近年来,随着一些新的研究手段包括化学分析、遗传工程、分子生物学、免疫学等方法的运用,推进了联合固氮领域的研究深度。综述了近年来发现的联合固氮菌的种类;联合固氮体系的形成过程（趋化、结合和侵入）;影响联合固氮的主要因素：自然条件、土著微生物的竞争、植物基因型差异和环境条件的变化、结合态氮（氨、亚硝酸盐、硝酸盐等）和氧。并从固氮联合作用测定方法、联合固氮菌的资源筛选、联合固氮工程菌的研究、联合固氮分子生态学研究方法4个方面,论述了联合固氮的研究现状。     

联合固氮

1定义氮素是构成生命物质最重要的元素之一。自然界只有某些微生物能直接将大气中的氮透过固氮酶还原成NH,这类微生物包括细菌,蓝绿藻、放线菌等。在固氮的细菌中有一类属于自由生活的类群,它们定殖于植物根表（有的能侵人根表皮和外皮层的细胞间隙）和近根土壤中,靠根系分泌物生存,繁延,与植物根系有密切的关系。但宿主植物并不形成特异分化的结构。植物与细菌之间的这种共生关系称联合共生固氮。这类固氮菌称联合固氮菌。联合共生固氮的概念是1976年由巴西Dobereiner实验室提出的。ZO年来联合固氮的研究受到世界各国科学家的重视…     

生物固氮及在可持续农业中的应用

氮是限制农业生产的重要营养元素.生物固氮指某些原核生物能利用体内的固氮酶将空气中的氮气还原为氨,为植物生长提供氮素.自然界中存在多种具有固氮能力的微生物,依据其固氮方式分为自生固氮、共生固氮和联合固氮三种类型.联合固氮茵通过趋化定殖在植物根表,并生长、固氮.     

内蒙古典型草原的生物量与生产力

氮素是植物生长和作物高产的限制因素。氮素来源主要是生物固氮和化学固氮。生物固氮是地球表面氮素的主要来源。在生物固氮中,目前最有效的还是豆科根瘤菌固氮。但近年来,禾本科植物根系与固氮菌的联合固氮作用引起了人们的广泛重视。联合固氮是自生固氮和共生固氮体系的中间类型,固氮细菌与相应联合植物之间具有较密切的相互影响,但又不象形成根瘤那样具有共生结构。这种联合固氮作用在自然界中广泛存在,各种作物,热带和亚热带牧草的根际和根表均有联合固氮菌存在,它们能提供土壤氮素。经测定水稻根际每个生长季非藻类的生物固氮量达25—30     

非豆科固氮树种-沙棘与微生物联合共生体的纯培养研究

Ｆｒａｎｋｉａ 非豆科树木共生固氮体系是自然界中重要的固氮生物资源 ,具有与Ｒｈｉｚｏｂｉｕｍ 豆科植物相似的固氮能力 ,在自然界能量循环和生态平衡中起着重要作用 ,而且与形成共生固氮的非豆科植物其抗逆性强 ,耐干旱、盐碱 ,抗高寒、瘠薄 ,能在一般豆科植物不能生长的环境下正常生长结瘤固氮。非豆科固氮树种 沙棘是三北地区重要垦荒先锋树种1) ,其果实富含多种维生素 ,具有重要的经济开发价值。同时 ,沙棘又是具有内生菌根真菌的固氮树种[1] 。本文通过对沙棘菌根、根瘤联合共生体人工构建技术以及沙棘联合共生的增效作用的研究 ,…     

内蒙古锡林郭勒草原不同固氮植物的固氮酶活性季节变化

 本文对内蒙古锡林郭勒典型草原栗钙土地带8种植物根瘤固氮酶活性的季节变化进行了测定。结果表明：1．不同种植物固氮酶活性表现了明显的季节性变化,夏季最高,平均值达到529.6n·mol C2H4／min·g,秋季较低,而春季只有夏季的10%左右。2．不同种植物,其根瘤固氮酶活性变化有不同的特点,杂花苜蓿是8种植物中固氮酶活性一直最高的,而小叶锦鸡儿与沙棘在8种植物中根瘤固氮酶活性一直较低,扁蓿豆,沙打旺前期较低,后期相对较高。3．同种植物在不同季节固氮酶活性表现了明显差别。     

非共生固氮菌及其固氮作用

生物固氮作用是生态系统中重要的氮素来源,参与固氮作用的微生物对植物的生长发育至关重要。与共生固氮微生物相比,非共生固氮微生物地域分布更广泛、种类更多,对全球生态系统中氮素循环有着重要意义。本文总结了非共生固氮菌的分类及系统发育,非共生固氮菌的群落构建过程和机制;归纳了不同生态系统(如草原、森林、海洋、农田等)、植物不同部位(如林冠、叶际、根际、根内、凋落物等)和土壤中非共生固氮菌的群落组成及固氮潜力的差异,以及影响非共生固氮菌群落组成和固氮潜力的主要因素(如气候因素、土壤理化性质、人为措施等);并整理了常用的研究非共生固氮菌及其固氮潜力的检测方法。     

铜对大叶相思-根瘤菌共生固氮体系的影响

报道了两种根瘤菌 (大叶相思、美丽胡枝子 )对Cu2 + 的耐受性以及植物 根瘤菌共生固氮体系在Cu2 + 胁迫下结瘤、固氮和生长的变化 ,讨论了大叶相思在矿山尾矿废弃地作为先锋植物结瘤固氮的可能性 .结果表明 ,大叶相思根瘤菌对Cu2 + 离子的耐受性较强 ,可以耐受Cu2 + <0 80mmol的离子浓度 ,Cu2 +对它的半致死浓度为 0 12 9mmol.在无菌砂培无重金属影响条件下 ,其固氮酶活性为 2 7C2 H4 ·μg·g-1·h-1,当Cu2 + >0 12 5mmol会导致大叶相思固氮酶活性急剧下降 ,其有效半抑制浓度 (EC50 )为0 15 1mmol,Cu2 + 为 0 5 0mmol完色抑制大叶相思固氮酶活性 ,不阻碍结瘤 ,但严重抑制植物生长发育 ,引起植物叶片白化、植株矮化 .在外加N源不接菌和不加N源接菌两种处理组间 ,Cu2 + <0 12 5mmol时 ,以不加N接菌处理对大叶相思生长有利 .大叶相思对Cu2 + 吸收积累根部高于地上组织 .     

VA菌根真菌、根瘤菌双接种对大豆增产效果的影响

本文论述了在自然状态下的土壤里,VA菌根真菌、根瘤菌双接种对大豆结瘤、菌根真菌侵染率、固氮酶活性及植株养分吸收状况的影响.研究结果表明,VA菌根真菌、根瘤菌双接种明显提高大豆结瘤能力与固氮酶活性,并可促进植物对养分的吸收,双接种比单接种VA菌根真菌处理提高产量140%,比单接种根瘤菌处理提高55.84%.     

土壤pH调控固氮植物和非固氮植物间的氮转移

氮作为构成蛋白质的主要成分, 是植物生长的必要营养物质。陆地生态系统普遍存在土壤氮缺乏的现象, 混交种植模式中固氮植物可以将生物固定的氮转移给非固氮植物, 是满足非固氮植物氮需求的途径之一。明确固氮和非固氮植物间氮转移的影响因素有助于恢复退化生态系统, 构建稳定群落, 增加生态系统生产力。为了量化环境及生物等因素对氮转移的影响, 该研究采用文献调研法, 对118组氮转移比例(氮转移量占非固氮植物氮含量的比值, P_transfer)文献和实验数据(包括21种固氮植物和23种非固氮植物)进行了线性混合模型分析。结果表明土壤pH是影响P_transfer变化的最主要因素(解释量为44.04%), 其次为年平均温度(解释量为9.14%)以及固氮与非固氮植物生物量比值(解释量为2.95%), 而作为随机因素的固氮和非固氮植物物种差异的解释量为16.52%。此外, 碱性土壤中P_transferr显著高于酸性土壤。在酸性土壤中, 年平均温度(解释量为12.49%)和土壤总氮含量(解释量为11.72%)是影响P_transfer差异的主要因素, P_transfer随着年平均温度和土壤总氮含量的增加而显著增加。而在碱性土壤中, P_transfer差异主要受到固氮与非固氮植物生物量比值(解释量为13.29%)、年降水量(解释量为10.73%)和土壤总氮含量(解释量为9.33%)的调控。相对于酸性土壤, 碱性土壤能够显著增加固氮与非固氮植物生物量比值进而增加P_transfer。同时, 在碱性土壤中P_transfer与年降水量和土壤总氮含量呈显著正相关关系。这些结果对提高固氮和非固氮植物间的氮转移, 有效缓解土壤氮对非固氮植物生长的限制以及构建稳定群落具有重要意义。     

Arbuscular mycorrhizal fungi can contribute to maintain antioxidant and carbon metabolism in nodules of Anthyllis cytisoides L. subjected to drought

The symbiosis legume-arbuscular mycorrhizal fungi-nitrogen fixing bacteria is of relevant interest in Mediterranean regions where Anthyllis cytisoides L. grows. In these areas, nitrogen is one of the nutrients that most limits plant growth. In addition, the long periods of water deficit decrease the diffusion rate of phosphorus and, consequently, also decrease the biological nitrogen fixation. It is well known that mycorrhizal fungi can improve phosphorus uptake and, recently, some authors have found that antioxidant activities in mycorrhizal plants can delay drought-induced nodule senescence. The objective of our work was to evaluate weather mycorrhizal fungi could preserve the nodule metabolism in A. cytisoides subjected to drought. Results showed that a low soil water content associated with an enhancement of soil compaction accelerated the senescence of nodules in both non-mycorrhizal and mycorrhizal plants. However, while total soluble protein, leghaemoglobin (Lb) content, as well as carbon and antioxidant metabolism significantly decreased in nodules from non-mycorrhizal A. cytisoides subjected to drought, nodules from stressed mycorrhizal plants maintained Lb levels, showed greater rates of carbon metabolism, and exhibited higher enzymatic activities related to the removal of reactive oxygen species. In addition to the greater activity of antioxidant enzymes, other mechanisms related or unrelated to enhanced nodule water status could also be implied in the better nodule functioning observed in mycorrhizal plants under stressful conditions.     

菌根真菌促进植物吸收利用氮素机制的研究进展

作为自然界最为普遍的一种植物共生体,菌根能够极大地促进植物对氮素的吸收和利用,其中菌根真菌在共生结构功能中发挥了重要作用。本文分别从菌根解剖构造、生理生化和分子生物学方面系统总结了菌根真菌促进植物吸收和利用氮素的研究现状。重点介绍了菌根真菌可利用的氮素形态及影响其利用的主要因素、菌根真菌的氮代谢途径GS-GOGAT以及菌根真菌中存在的鸟氨酸循环途径,指出精氨酸是菌丝内氮转运的主要形式,NH3可能为菌根真菌和植物界面质外体的主要转运形式。     

Changes in alfalfa forage quality and stem carbohydrates induced by arbuscular mycorrhizal fungi and elevated atmospheric CO2

Alfalfa is a widely distributed forage legume whose leaves are high in protein content and whose stems are suitable for bioethanol production. However, alfalfa forage digestibility, quality and yield may vary under future climate change scenarios. This legume can establish double symbiosis with nitrogen‐fixing bacteria and arbuscular mycorrhizal fungi (AMF). The presence of AMF can modify the evolution of biomass production and partitioning during the vegetative growth of alfalfa. We hypothesised that mycorrhizal symbiosis may change the quantity and/or quality of carbohydrates and lignin in leaves and/or stems of alfalfa, with these changes being dependent on the atmospheric CO₂ concentration at which plants are grown. Results showed that mycorrhizal alfalfa plants exposed to elevated CO₂ had improved leaf, stem and root biomass, enhanced amount of hemicellulose and decreased concentration of lignin in cell walls of leaves as well as increased levels of glucose and fructose in stems compared with non‐mycorrhizal alfalfa. These results indicated improved forage quality (leaves) and enhanced potential for bioethanol conversion (stems) in mycorrhizal alfalfa cultivated under elevated CO₂. Moreover, the potential of stems for producing CH₄ reinforced their suitability for the conversion of biomass into bioethanol.     

Mycorrhizal benefits on native plants of the Caatinga,a Brazilian dry tropical forest

In the terrestrial ecosystems, soil is an important component, characterized by holding high diversity of microorganisms which play a key role for productivity and vegetal composition. The group of symbionts microorganisms stands out for contributing directly to the growth and plant nutrition, and among them, the arbuscular mycorrhizal fungi form one of the oldest and well established associations. In order to increase the knowledge and contribute for further research with AMF and plants of Caatinga, in this review we compile data from previous studies on the effects of symbiosis between arbuscular mycorrhizal fungi (AMF) and plants of the Caatinga, a type of dry tropical forest found in the northeast of Brazil. These studies collected data under various experimental conditions, emphasizing fungal efficiency and host responsiveness in soils with varied fertility. From our analysis we conclude that in general the symbiotic efficiency on these plants depends on many factors, such as the plant-fungi combination, fertility and soil type. Furthermore, in leguminosae the impact of a joint inoculation with nitrogen fixing bacteria must be taken into account. Claroideoglomus etunicatum was the most tested AMF species benefiting almost all plants tested. Approximately 30 plant species were studied regarding possible benefits provided by AMF and of these only Hymenea courbaril and Aspidosperma pyrifolium did not respond to mycorrhization. Higher efficiency of the mycorrhizal symbiosis can be obtained in soils with low P levels, emphasizing the essential role of these microorganisms in the growth and survival of plant species from the Caatinga biome.     

菌根真菌与植物共生营养交换机制研究进展

菌根是陆地生态系统普遍存在的、由土壤中的菌根真菌侵染宿主植物根系形成的联合共生体.菌根的建立是以共生体双方的营养交换为基础的:菌根真菌从土壤中吸收氮、磷等营养物质并转运给宿主植物,供其生长;作为交换,植物则以脂质或糖的形式向菌根真菌提供其生长所必需的碳水化合物.近年来,菌根真菌与宿主植物间的营养交换机制一直是研究的热点,国内外对菌根真菌介导的植物营养物质吸收和转运机制的研究也取得了巨大进展.本文综述了丛枝和外生两种菌根真菌与宿主植物间营养交换的最新研究进展,尤其是碳、氮、磷等几种重要营养物质的吸收与双向转运机制,以及营养交换在菌根形成中的潜在调控作用,并对目前存在的关键问题和未来研究方向进行了分析和展望,这对菌根模型的建立及菌根效益的优化具有重要意义.     

An anthocyanin marker for direct visualization of plant transformation and its use to study nitrogen-fixing nodule development

Journal of Plant Research - The development and functioning of the nitrogen fixing symbiosis between legume plants and soil bacteria collectively called rhizobia requires continuous chemical...     

Phosphorus and Nitrogen Regulate Arbuscular Mycorrhizal Symbiosis in Petunia hybrida

Phosphorus and nitrogen are essential nutrient elements that are needed by plants in large amounts. The arbuscular mycorrhizal symbiosis between plants and soil fungi improves phosphorus and nitrogen acquisition under limiting conditions. On the other hand, these nutrients influence root colonization by mycorrhizal fungi and symbiotic functioning. This represents a feedback mechanism that allows plants to control the fungal symbiont depending on nutrient requirements and supply. Elevated phosphorus supply has previously been shown to exert strong inhibition of arbuscular mycorrhizal development. Here, we address to what extent inhibition by phosphorus is influenced by other nutritional pathways in the interaction between Petunia hybrida and R. irregularis. We show that phosphorus and nitrogen are the major nutritional determinants of the interaction. Interestingly, the symbiosis-promoting effect of nitrogen starvation dominantly overruled the suppressive effect of high phosphorus nutrition onto arbuscular mycorrhiza, suggesting that plants promote the symbiosis as long as they are limited by one of the two major nutrients. Our results also show that in a given pair of symbiotic partners (Petunia hybrida and R. irregularis), the entire range from mutually symbiotic to parasitic can be observed depending on the nutritional conditions. Taken together, these results reveal complex nutritional feedback mechanisms in the control of root colonization by arbuscular mycorrhizal fungi.     

Plant Kin Recognition Enhances Abundance of Symbiotic Microbial Partner

Background

The stability of cooperative interactions among different species can be compromised by cheating. In the plant-mycorrhizal fungi symbiosis, a single mycorrhizal network may interact with many plants, providing the opportunity for individual plants to cheat by obtaining nutrients from the fungi without donating carbon. Here we determine whether kin selection may favour plant investment in the mycorrhizal network, reducing the incentive to cheat when relatives interact with a single network.

Methodology/Principal Findings

We show that mycorrhizal network size and root colonization were greater when Ambrosia artemisiifolia L. was grown with siblings compared to strangers. Soil fungal abundance was positively correlated with group leaf nitrogen, and increased root colonization was associated with a reduced number of pathogen-induced root lesions, indicating greater benefit to plants grown with siblings.

Conclusions/Significance

Plants can benefit their relatives through investment in mycorrhizal fungi, and kin selection in plants could promote the persistence of the mycorrhizal symbiosis.     

Mycorrhizal status of Cyperaceae from New Caledonian ultramafic soils: effects of phosphorus availability on arbuscular mycorrhizal colonization of Costularia comosa under field conditions

Plants from the Cyperaceae family (sedges), usually considered as non-mycorrhizal, constitute almost exclusively the herbaceous stratum of the ultramafic maquis in New Caledonia. These plants are pioneers and are important for the ecological restoration of mined areas. Costularia comosa, one of the most common sedges in this environment, was grown under field conditions on ultramafic soil, fertilized or not with phosphate and/or nitrogen. Results showed that the addition of phosphate to the soil induced a clear increase in mycorrhizal colonization of C. comosa and an increase in arbuscule abundance, reflecting the establishment of a functional mycorrhizal symbiosis. Significant positive correlations were found among mycorrhizal parameters and plant or soil phosphorus concentrations. Nitrogen fertilization did not affect mycorrhizal colonization of C. comosa. The improvement in mycorrhizal colonization by phosphate fertilization did not influence significantly nickel concentrations in the roots and shoots of plants. This study demonstrated that phosphate fertilization of ultramafic soil improved mycorrhizal colonization of C. comosa, with formation of a functional symbiosis under field conditions.     

丛枝菌根形成过程及其信号转导途径

丛枝菌根是由一类土壤中古老的丛枝菌根真菌与植物根系形成的互利互惠共生体。通过共生作用丛枝菌根真菌帮助宿主植物提高水和矿质营养（特别是磷）的吸收效率。作为回报,大约20%的光合作用产物被转移到丛枝菌根真菌中,供其完成自身的生活史。丛枝菌根形成的过程中,需要植物与丛枝菌根真菌之间进行一系列信号分子的识别、交换以及信号转导作用,这一过程由一系列植物和菌根真菌的基因控制。首先,植物会分泌一种植物激素——独角金内酯来诱导菌根真菌加速分支,而菌根真菌也会分泌脂质几丁寡糖促进植物与其形成菌根。加速分支的菌根真菌接触到植物根部以后,会附着在植物根的表皮并形成附着胞,通过附着胞穿透植物根的表皮,最后进入维管组织附近的皮层细胞并在其中不断进行二叉分支,形成特有的丛枝结构。通过对模式植物共生现象的研究,已经发现很多植物基因参与到共生形成的信号转导过程中,包括早期植物反应的基因、菌根与根瘤共生共同需要的转导因子以及菌根特异的信号分子等。本文对菌根的形成过程及信号转导途径进行详细的介绍,为人们深入研究菌根关系提供参考。