[基于细菌从菌根真菌和土壤向植物组织迁移的固氮共生进化]。

Pub Date : 2016-09-01
N A Provorov, O Yu Shtark, E A Dolgikh
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引用次数: 0

摘要

提出了固氮植物共生体从土壤重氮营养体和形成丛枝菌根(AM)的肾小球真菌卫星中出现的假说。这种普遍形式的植物-微生物共生可能是由祖先陆地植物(流根植物、精神植物)和am真菌(吸收土壤磷酸盐)和细菌(固定大气CO2和/或N2)组成的微生物联合体整合而成的。这些细菌从am -真菌菌丝中释放到植物组织中,引发了能够在植物中进行真菌独立繁殖的基因型的选择,并在细菌基因组中固定了用于合成刺激共生结构发展的几丁质样信号因子的基因。这种进化的早期阶段可能以形成固定n2的合胞菌为代表,后期阶段-由来自Eurosid I分支的dicotes与根瘤菌(α-和β-变形菌)和放线菌Frankia形成结节状共生。这些共生关系的出现可能是基于土壤和内生细菌向储存器官(改良茎或侧根)的迁移,在那里不仅为固氮创造了最佳条件,而且为细菌向增加共生活性的进化创造了最佳条件。这种进化导致了原生根瘤菌(慢生根瘤菌,伯克霍尔德菌)的出现,这些根瘤菌作为共生基因的供体,为广谱的微生物转化为次生根瘤菌(根瘤菌,中华根瘤菌)。结核共生的后续进化旨在提高寄主植物共生氮营养的效率,并遵循以下两种情况:(i)“昂贵”,通过将细菌转化为不可复制的类细菌来增加固氮活性;(ii)“经济”,基于获得确定的结核结构和尿素氮同化。
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[Evolution of nitrogen-fixing symbioses based on the migration of bacteria from mycorrhizal fungi and soil into the plant tissues].

The hypothesis is suggested on the emergence of N2-fixing plant symbionts from soil diazotrophs and from the satellites of Glomeromycota fungi forming arbuscular mycorrhizae (AM). This universal form of plant-microbe symbiosis possibly appeared from integration of ancestral land plants (rhyniophytes, psylophytes) and microbial consortia composed of AM-fungi assimilating soil phosphates and bacteria fixing atmospheric CO2 and/or N2. Releasing of these bacteria from AM-fungal hyphae into the plant tissues elicited the selection of genotypes capable of the fungi-independent multiplication in planta, as well as the fixation in bacterial genomes of the genes for synthesis of chitin-like signal factors stimulating the development of symbiotic structures. An early stage of this evolution might been represented by formation of N2-fixing syncyanoses, the late stage - by formation of nodular symbioses of dicots from Eurosid I clade with rhizobia (α- and β-proteobacteria) and with actinobacteria Frankia. Emergence of these symbioses was possibly based on the migration of soil and endophytic bacteria into the storage organs (modified stems or lateral roots), where the optimal conditions were established not only for N2 fixation but also for the evolution of bacteria towards an increased symbiotic activity. This evolution resulted in the emergence of primary rhizobia (Bradyrhizobium, Burkholderia) which acted as the donors of sym-genes for a broad spectrum of microbes transformed into the secondary rhizobia (Rhizobium, Sinorhizobium). The succeeding evolution of nodular symbioses was directed at an increased efficiency of symbiotrophic nitrogen nutrition in host plants following two scenarios: (i) “expensive”, based on the increase of N2- fixing activity via transformation of bacteria into non-reproducible bacteroids; (ii) “economic”, based on acquiring the determinate nodule structure and ureide nitrogen assimilation.

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