出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2016/03/19 06:42:24」(JST)
菌根（きんこん）は、菌類が植物の根に侵入して形成する特有の構造を持った共生体。菌根を作る菌類を菌根菌という。
菌根には7つの主要なタイプがあり、それぞれ関与する菌類や植物が異なり、構造も異なる。アーバスキュラー菌根(かつてはVA菌根と呼んだ)、外生菌根または外菌根、内外生菌根、アルブトイド菌根、モノトロポイド菌根、エリコイド菌根、ラン菌根の７つの主要なタイプと、最近認識されてきたものにハルシメジ型菌根がある。
菌根はよく細菌と植物の根との共生体である根粒と混同されるが、菌根の共生微生物は真菌であり、宿主植物が7タイプあわせると陸上植物の大半といえるほど多く、窒素固定を行わないなど、根粒とは全く異なるものである。一方で、近年のアーバスキュラー菌根の形成に関する研究から、菌根と根粒の形成過程に関与する植物側の遺伝子には共通するものも多いことが明らかになっている。
菌根の主要な機能としては、一般に土壌中の栄養塩類、すなわち肥料分の吸収と宿主への輸送、土壌病害への抵抗性の向上、水分吸収能力の強化の3点が挙げられる。これに対し植物が菌根菌に光合成産物(エネルギー)を与えるという相利共生を営んでいるとされるが、これには例外も多い。アーバスキュラー菌根や外菌根ではこの相利共生が成立するものも多いが、たとえばホンゴウソウ科やヒナノシャクジョウ科などの無葉緑植物もアーバスキュラー菌根を形成する。共生相手が無葉緑植物では菌根菌は光合成産物を得ることはできず、アーバスキュラー菌根菌は絶対共生者で腐生的に養分獲得を行うこともできないが、この場合は同一の菌糸体が他方で光合成を行う緑色植物とも共生関係を結んでおり、そこから光合成産物を得てその一部を無葉緑植物に渡していると考えられている。そのため、エネルギー的にはホンゴウソウ科やヒナノシャクジョウ科の植物は菌に寄生していることになり、菌従属栄養植物と呼ばれている。
アルブトイド菌根、モノトロポイド菌根、ラン菌根では基本的に植物が菌に寄生する関係となっており、モノトロポイド菌根を形成する無葉緑植物のギンリョウソウやラン菌根を形成するオニノヤガラやツチアケビなどの無葉緑ランもまた菌従属栄養植物である。アルブトイド菌根を形成するイチヤクソウ類も強く菌根菌に依存した生活様式をもっている。かつては他の植物に寄生しない無葉緑植物は土壌中の腐植などから養分を獲得していると想像され腐生植物と呼ばれたが、近年それらは菌根から養分を獲得しておりその起源も必ずしも腐植とは限らないことが明らかになってきたため、菌従属栄養植物という言葉が使われるようになってきた。
| この項目は、菌類に関連した書きかけの項目です。この項目を加筆・訂正などしてくださる協力者を求めています（P:生き物と自然／PJ生物）。 |
| It has been suggested that Mycorrhizal networks be merged into this article. (Discuss) Proposed since April 2013. |
A mycorrhiza (Greek: μυκός, mykós, "fungus", and ρίζα, riza, "root",[1] pl. mycorrhizae or mycorrhizas) is a symbiotic association composed of a fungus and roots of a vascular plant.[2] In a mycorrhizal association, the fungus colonizes the host plant's roots, either intracellularly as in arbuscular mycorrhizal fungi (AMF or AM), or extracellularly as in ectomycorrhizal fungi. They are an important component of soil life and soil chemistry. The association is generally mutualistic, but occasionally weakly pathogenic.[clarification needed]
Fungi in mycorrhizae form a mutualistic relationship with the roots of most plant species. The roots in the relationship, and the plants themselves are referred to as mycorrhizal if mycorrhizae are formed. While only a small proportion of all species has been examined, 95% of those plant families[which?] are predominantly[clarification needed] mycorrhizal.[3] They are named after their presence in the plant's rhizosphere (root system).
Recent research with ectomycorrhizal plants in boreal forests has indicated that mycorrhizal fungi and plants have a relationship that may be more complex than simply mutualistic. This relationship was noted when mycorrhizal fungi were unexpectedly found hoarding nitrogen from plant roots in times of nitrogen scarcity. Researchers[who?] argue that some mycorrhizae distribute nutrients based upon the environment with surrounding plants and other mycorrhizae. Researchers go on to explain how this updated model explains why mycorrhizae do not alleviate plant nitrogen limitation, and why plants can switch abruptly from a mixed strategy with both mycorrhizal and nonmycorrhizal roots to a purely mycorrhizal strategy as soil nitrogen availability declines.[4]
This mutualistic association provides the fungus with relatively constant and direct access to carbohydrates, such as glucose and sucrose.[5] The carbohydrates are translocated from their source (usually leaves) to root tissue and on to the plant's fungal partners. In return, the plant gains the benefits of the mycelium's higher absorptive capacity for water and mineral nutrients due to the large surface area of fungal hyphae, which are much finer than plant roots, thus improving the plant's mineral absorption capabilities.[6]
Plant roots alone may be incapable of taking up phosphate ions that are demineralized in soils with a basic pH. The mycelium of the mycorrhizal fungus can, however, access these phosphorus sources, and make them available to the plants they colonize.[7] Thus many plants are able to obtain phosphate, without using soil as a source. For example, in some dystrophic forests, large amounts of phosphate are taken up by mycorrhizal hyphae acting directly on leaf litter, bypassing the need for soil uptake.[8] Inga alley cropping, proposed as an alternative to slash and burn rainforest destruction,[9] relies upon mycorrhiza within the Inga Tree root system to prevent the rain from washing phosphorus out of the soil.[10] In some cases, the transport of water, carbon, and nutrients could be done directly from plant to plant through mycorrhizal networks that are underground hyphal networks created by mycorrhizal fungi that connect individual plants together.[11]
Suillus tomentosus, a basidiomycete fungus, produces specialized structures known as tuberculate ectomycorrhizae with its plant host lodgepole pine (Pinus contorta var. latifolia). These structures have been shown to host nitrogen fixing bacteria which contribute a significant amount of nitrogen and allow the pines to colonize nutrient-poor sites.[12]
The mechanisms of increased absorption[clarification needed] are both physical and chemical. Mycorrhizal mycelia are much smaller in diameter than the smallest root, and thus can explore a greater volume of soil, providing a larger surface area for absorption. Also, the cell membrane chemistry of fungi is different from that of plants (including organic acid excretion which aids in ion displacement[13]). Mycorrhizas are especially beneficial for the plant partner in nutrient-poor soils.[14]
Mycorrhizal plants are often more resistant to diseases, such as those caused by microbial soil-borne pathogens. AMF was also significantly correlated with soil biological fertility variables such as soil fungi and soil bacteria, including soil disease.[citation needed] Furthermore, AMF was significantly correlated with soil physical variable, but only with water level and not with aggregate stability.[15][16] and are also more resistant to the effects of drought.[17][18][19] It is known the significance of arbuscular mycorrhizal fungi alleviation of salt stress and their beneficial effects on plant growth and productivity. Although salinity can affect negatively arbuscular mycorrhizal fungi, many reports show improved growth and performance of mycorrhizal plants under salt stress conditions [20]
Plants grown in sterile soils and growth media often perform poorly without the addition of spores or hyphae of mycorrhizal fungi to colonise the plant roots and aid in the uptake of soil mineral nutrients.[21] The absence of mycorrhizal fungi can also slow plant growth in early succession or on degraded landscapes.[22] The introduction of alien mycorrhizal plants to nutrient-deficient ecosystems puts indigenous non-mycorrhizal plants at a competitive disadvantage.[23]
Fungi have been found to have a protective role for plants rooted in soils with high metal concentrations, such as acidic and contaminated soils. Pine trees inoculated with Pisolithus tinctorius planted in several contaminated sites displayed high tolerance to the prevailing contaminant, survivorship and growth.[24] One study discovered the existence of Suillus luteus strains with varying tolerance of zinc. Another study discovered that zinc-tolerant strains of Suillus bovinus conferred resistance to plants of Pinus sylvestris. This was probably due to binding of the metal to the extramatricial mycelium of the fungus, without affecting the exchange of beneficial substances.[23]
At around 400 million years old, the Rhynie chert contains an assemblage of fossil plants preserved in sufficient detail that mycorrhizas have been observed in the stems of Aglaophyton major.[25]
Mycorrhizas are present in 92% of plant families studied (80% of species),[26] with arbuscular mycorrhizas being the ancestral and predominant form,[26] and the most prevalent symbiotic association found in the plant kingdom.[5] The structure of arbuscular mycorrhizas has been highly conserved since their first appearance in the fossil record,[25] with both the development of ectomycorrhizas, and the loss of mycorrhizas, evolving convergently on multiple occasions.[26]
Mycorrhizas are commonly divided into ectomycorrhizas and endomycorrhizas. The two types are differentiated by the fact that the hyphae of ectomycorrhizal fungi do not penetrate individual cells within the root, while the hyphae of endomycorrhizal fungi penetrate the cell wall and invaginate the cell membrane.[27][28] Endomycorrhiza includes arbuscular, ericoid, and orchid mycorrhiza, while arbutoid mycorrhizas can be classified as ectoendomycorrhizas. Monotropoid mycorrhizas form a special category.
Endomycorrhizas are variable and have been further classified as arbuscular, ericoid, arbutoid, monotropoid, and orchid mycorrhizas.[29] Arbuscular mycorrhizas, or AM (formerly known as vesicular-arbuscular mycorrhizas, or VAM), are mycorrhizas whose hyphae enter into the plant cells, producing structures that are either balloon-like (vesicles) or dichotomously branching invaginations (arbuscules). The fungal hyphae do not in fact penetrate the protoplast (i.e. the interior of the cell), but invaginate the cell membrane. The structure of the arbuscules greatly increases the contact surface area between the hypha and the cell cytoplasm to facilitate the transfer of nutrients between them.
Arbuscular mycorrhizas are formed only by fungi in the division Glomeromycota. Fossil evidence[25] and DNA sequence analysis[30] suggest that this mutualism appeared 400-460 million years ago, when the first plants were colonizing land. Arbuscular mycorrhizas are found in 85% of all plant families, and occur in many crop species.[26] The hyphae of arbuscular mycorrhizal fungi produce the glycoprotein glomalin, which may be one of the major stores of carbon in the soil. Arbuscular mycorrhizal fungi have (possibly) been asexual for many millions of years and, unusually, individuals can contain many genetically different nuclei (a phenomenon called heterokaryosis).[31]
Ectomycorrhizas, or EcM, are typically formed between the roots of around 10% of plant families, mostly woody plants including the birch, dipterocarp, eucalyptus, oak, pine, and rose[26] families, orchids,[32] and fungi belonging to the Basidiomycota, Ascomycota, and Zygomycota. Some EcM fungi, such as many Leccinum and Suillus, are symbiotic with only one particular genus of plant, while other fungi, such as the Amanita, are generalists that form mycorrhizas with many different plants.[33] An individual tree may have 15 or more different fungal EcM partners at one time.[34] Thousands of ectomycorrhizal fungal species exist, hosted in over 200 genera. A recent study has conservatively estimated global ectomycorrhizal fungal species richness at approximately 7750 species, although, on the basis of estimates of knowns and unknowns in macromycete diversity, a final estimate of ECM species richness would probably be between 20000 and 25000.[35]
Ectomycorrhizas consist of a hyphal sheath, or mantle, covering the root tip and a Hartig net of hyphae surrounding the plant cells within the root cortex. In some cases the hyphae may also penetrate the plant cells, in which case the mycorrhiza is called an ectendomycorrhiza. Outside the root, Ectomycorrhizal extramatrical mycelium forms an extensive network within the soil and leaf litter.
Nutrients can be shown to move between different plants through the fungal network. Carbon has been shown to move from paper birch trees into Douglas-fir trees thereby promoting succession in ecosystems.[36] The ectomycorrhizal fungus Laccaria bicolor has been found to lure and kill springtails to obtain nitrogen, some of which may then be transferred to the mycorrhizal host plant. In a study by Klironomos and Hart, Eastern White Pine inoculated with L. bicolor was able to derive up to 25% of its nitrogen from springtails.[37][38]
The first genomic sequence for a representative of symbiotic fungi, the ectomycorrhizal basidiomycete Laccaria bicolor, has been published.[39] An expansion of several multigene families occurred in this fungus, suggesting that adaptation to symbiosis proceeded by gene duplication. Within lineage-specific genes those coding for symbiosis-regulated secreted proteins showed an up-regulated expression in ectomycorrhizal root tips suggesting a role in the partner communication. Laccaria bicolor is lacking enzymes involved in the degradation of plant cell wall components (cellulose, hemicellulose, pectins and pectates), preventing the symbiont from degrading host cells during the root colonisation. By contrast, Laccaria bicolor possesses expanded multigene families associated with hydrolysis of bacterial and microfauna polysaccharides and proteins. This genome analysis revealed the dual saprotrophic and biotrophic lifestyle of the mycorrhizal fungus that enables it to grow within both soil and living plant roots.
Ericoid mycorrhizas are the third of the three more ecologically important types. They have a simple intraradical (grow in cells) phase, consisting of dense coils of hyphae in the outermost layer of root cells. There is no periradical phase and the extraradical phase consists of sparse hyphae that don't extend very far into the surrounding soil. They might form sporocarps (probably in the form of small cups), but their reproductive biology is little understood.[28]
Ericoid mycorrhizas have also been shown to have considerable saprotrophic capabilities, which would enable plants to receive nutrients from not-yet-decomposed materials via the decomposing actions of their ericoid partners.[41]
This type of mycorrhiza involves plants of the Ericaceae subfamily Arbutoideae. It is however different from ericoid mycorrhiza and resembles ectomycorrhiza, both functionally and in terms of the fungi involved.[citation needed] The difference to ectomycorrhiza is that some hyphae actually penetrate into the root cells, making this type of mycorrhiza an ectendomycorrhiza.[citation needed]
This type of mycorrhiza occurs in the subfamily Monotropoideae of the Ericaceae. These plants are heterotrophic or mixotrophic and derive their carbon from the fungus partner. This is thus a non-mutualistic, parasitic type of mycorrhizal symbiosis.[citation needed]
All orchids are myco-heterotrophic at some stage during their lifecycle and form orchid mycorrhizas with a range of basidiomycete fungi.[citation needed] Their hyphae penetrate into the root cells and form typical coils.[citation needed]
Associations of fungi with the roots of plants have been known since at least the mid-19th century. However early observers simply recorded the fact without investigating the relationships between the two organisms.[42] This symbiosis was studied and described by Franciszek Kamieński in 1879–1882.[43] Further research was carried out by Albert Bernhard Frank, who introduced the term mycorrhiza in 1885.[44]
| Fungi portal |
| Wikisource has the text of the 1920 Encyclopedia Americana article Mycorriza. |
|
| リンク元 | 「菌根」 |
| 拡張検索 | 「mycorrhizal」「mycorrhizae」 |