Provorov N A
Zh Obshch Biol. 2009 Jan-Feb;70(1):10-34.
In spite of enormous taxonomic, structural and functional diversity of plant-microbe interactions, they are characterized by a historical succession which allows us to consider different forms of symbioses as the components of an evolutionary continuum. Their ancestral form is represented by arbuscular mycorrhiza (AM) which originated at the outset of terrestrial flora evolution and constituted a key factor for the land colonization by plants. In the course of AM evolution the plant acquired a basal set of genes for regulating the performance of microbes which colonize the root tissues. Later, these genes were repeatedly reorganized to meet the involvement of novel mutualistic symbionts (N2-fixing bacteria, ectomycorrhizal fungi, endophytes and epiphytes) and pathogens into the symbiotic interactions. Form the microbial side, the evolutionary succession of mutualism and antagonism is restricted to the defensive symbioses formed by plants with the ergot fungi, Clavibacter, Bacillus and Pseudomonas bacteria. Involvement of the similar systems for symbiotic interactions may be related to convergent evolution in the distant microorganisms (adaptation to the conservative host defense/regulatory factors), to molecular mimicry (imitation of the mechanisms of interaction used by the more ancient symbionts) or to the horizontal gene transfer. The hypotheses of the successive substitution of symbionts is suggested to address the relationships between AM and N2-fixing nodular symbioses in dicotyledons plants. AM formation is considered as a source of preadaptations responsible for the substitution of glomalean fungi which occupied the plant symbiotic compartments by the actinomycetes Frankia (in Rosid I plants) which were exchanged for the more competitive root nodule bacteria (in legumes). The development of nutritional symbioses with microbes is considered as an ancestral function of plant roots which were later supplemented or substituted with the function of assimilating the soil nutrients.
尽管植物与微生物相互作用在分类学、结构和功能上具有巨大的多样性,但它们具有历史连续性,这使我们能够将不同形式的共生关系视为进化连续体的组成部分。它们的原始形式以丛枝菌根(AM)为代表,其起源于陆地植物群进化之初,是植物在陆地定殖的关键因素。在AM进化过程中,植物获得了一组基础基因,用于调节定殖于根组织的微生物的性能。后来,这些基因不断重组,以适应新的互利共生体(固氮细菌、外生菌根真菌、内生菌和附生植物)和病原体参与共生相互作用。从微生物方面来看,共生和拮抗的进化连续性仅限于植物与麦角菌、棒状杆菌、芽孢杆菌和假单胞菌形成的防御性共生关系。类似的共生相互作用系统的参与可能与远距离微生物的趋同进化(适应保守的宿主防御/调节因子)、分子模拟(模仿更古老共生体使用的相互作用机制)或水平基因转移有关。共生体连续替代的假说是为了解决双子叶植物中AM与固氮根瘤共生关系之间的联系而提出的。AM的形成被认为是一种预适应的来源,它导致占据植物共生区室的球囊霉真菌被放线菌弗兰克氏菌(在蔷薇I类植物中)取代,而后者又被更具竞争力的根瘤菌(在豆科植物中)所替代。与微生物形成营养共生关系的发展被认为是植物根系的一种原始功能,后来这种功能被吸收土壤养分的功能所补充或替代。