Hause,B. and Fester,T.(2005): Molecular and cell biology of arbuscular mycorrhizal symbiosis. Planta, 221, 184-196.

アーバスキュラー菌根共生の分子・細胞生物学』


Abstract
 The roots of most extant plants are able to become engaged in an interaction with a small group of fungi of the fungal order Glomales (Glomeromycota). This interaction - arbuscular mycorrhizal (AM) symbiosis - is the evolutionary precursor of most other mutualistic root-microbe associations. The molecular analysis of this interaction can elucidate basic principles regarding such associations. This review summarizes our present knowledge about cellular and molecular aspects of AM. Emphasis is placed on morphological changes in colonized cells, transfer of nutrients between both interacting partners, and plant defence responses. Similarities to and differences from other associations of plant and microorganisms are highlighted regarding defence reactions and signal perception.

Keywords: Arbuscular mycorrhizal fungi; Defence response; Induced systemic resistance; Morphology of arbuscule-containing cells; Nutrient transfer; Signal transduction』

要旨
 ほとんどの現存する植物の根は、接合菌類のグロムス目(Glomeromycota)という菌類の小さなグループと相互作用で結びつくことができる。この相互作用−アーバスキュラー菌根(AM)共生−は、ほとんどの他の相利共生的な根−微生物群集の進化的な先駆にあたる。この相互作用を分子レベルで解析すると、そのような群集に関する基本的な原理を解明できる。このレビューは、AMの細胞と分子の面から現在の知識をまとめている。コロニーが形成された細胞の形態変化、相互作用を行っている両者間の栄養の輸送、および植物の防御反応に主眼がおかれている。植物と微生物の他の群集との類似点および相違点が、防御反応と信号認知に関して強調されている。』

Introduction
The fungal partner
Model plants in AM research
Cytological features of AM plant roots
Transfer of nutrients between plants and fungi
Defence reactions of plant roots during colonization by AM fungi
Induced systemic resistance in AM plants
Signalling pathways in AM
Conclusions/perspectives
Acknowledgements
References

Fig. 1 Survey of the morphology of an arbuscule-containing root cortex cell.
a
Scheme of a young arbuscule (red) within a root cortex cell. The fungal hypha penetrates the cell wall (grey) and undergoes branching leading to the formation of an arbuscule. The arbuscule is surrounded by the plant cytoplasm (orange) which contains high numbers of organelles (plastids - dark green, mitochondria - purple, ER - yellow). Fungal hyphae and plant cytoplasm are separated by the periarbuscular membrane (light grey). The plant cell nucleus (blue) moves into the centre of the arbuscule; the vacuole (light green) fragments after the arbuscule is fully developed.
b GFP-labelled plastids forming a network-like structure, which covers the arbuscule. Mycorrhizal roots of stably transformed tobacco plants, expressing a plastid-directed GFP (kindly provided by M. Hanson, New York, USA), were analysed by confocal laser scanning microscopy (CLSM). The superposition of 20 optical sections is shown. Bar represents 10 μm.
c Visualization of ER in arbusculated cells. Mycorrhizal roots of transgenic N. benthamiana plants expressing ER-targeted GFP (kindly provided by D. Baulcombe, Norwich) were processed for immunolocalization according to Hans et al. (2004). Cross-sections were probed with anti-GFP antibody followed by a fluorescence-labelled secondary antibody. Micrographs were taken by CLSM showing GFP (green), DAPI-stained nuclei (blue) and fungal structures stained with WGA-TRITC (red). The superposition of 28 optical sections is shown. Bar represents 10 μm.

arbuscule=樹枝状体、cortex=皮層、hypha=菌糸、cell wall=細胞壁、cytoplasm=細胞質、organelle=細胞小器官、plastid=色素体、mitochondrion(複数はmitochondria)=ミトコンドリア、ER=小胞体、membrane=膜、nucleus=、vacuole=液胞、GFP(Green Fluorescent Protein)〔蛍光蛋白質〕、CLSM=共焦点レーザースキャン顕微鏡法、transgenic=遺伝形質転換性、antibody=抗体、DAPI(4',6-Diamidino-2-phenylindole)〔青紫色蛍光色素〕、WGA-TRITC(wheat germ agglutinin(コムギ胚凝集素) conjugated to tetramethylrhodamine isothiocyanate-dextran〔赤橙色蛍光色素〕)。

Fig. 2 Nutrient transfer in AM roots.
Enzymes and transporters described to be specifically induced in AM roots are indicated. Membrane transport of most metabolites can be expected to be pH-dependent and to be powered by the activity of plant (1) and fungal (2) H+ -ATPases. Fungal H+ -ATPases have been described not to be restricted to arbuscules, suggesting active transport at intercellular hyphae as well. Sucrose from the phloem is either cleaved by apoplastic invertases and taken up by the plant (3) or fungal hexose transporters or imported into root cortical cells and cleaved there by a cytoplasmic sucrose synthase (4). The fungus transforms hexoses rapidly into trehalose, which is either metabolized by the pentose phosphate pathway, or used for the biosynthesis of glycogen and lipids. These compounds are then exported to fungal vesicles or to the external mycelium. The plant cell takes up phosphate from the periarbuscular space using specific, H+ -dependent plant phosphate transporters (5). Regarding nitrogen supply, AM-induced plant nitrate transporters (6) have been found, suggesting a similar transport mechanism as referring to phosphate. On the other hand, the observation of increased transcript levels of a fungal nitrate reductase (7) suggests the transfer of nitrogen in a reduced form (as ammonium or in an organic form). AA Amino acids.

enzyme=酵素、metabolite=代謝(産)物、ATPase=ATPアーゼ、arbuscule=樹枝状体、intercellular=細胞間、sucrose=スクロース、phloem=師部、invertase=インベルターゼ、hexose=ヘキソース、pentose phosphate pathway=ペントースリン酸経路、glycogen=グリコーゲン、lipid=脂質、vesicle=嚢状体、mycelium=菌糸体

〔Hause,B. and Fester,T.(2005): Molecular and cell biology of arbuscular mycorrhizal symbiosis. Planta, 221, 184-196.から〕


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