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The bi-directional movement of nutrients - carbon (C) from plant to fungus, and soil nutrients from fungus to plant - is the essential feature of a mycorrhiza, and is believed to be the basis for the mutualistic association. Aside from monotropoid mycorrhizas, the juvenile stages of all orchids, and the whole lives of a few orchid species (which are achlorophyllous) there is always a 2-way transfer of nutrients. |
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As a rule, mycorrhizal infection enhances plant growth, by increasing nutrient uptake in up to 3 ways: i) by increasing the surface area of absorption within the soil, ii) by mobilising sparingly available nutrient sources from unavailable compounds, and iii) by excreting chelating compounds or ectoenzymes (Marschner & Dell, 1994). |
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![]() The effects of mycorrhizal inoculation on crop growth. Right-hand block inoculated with AM fungi, left-hand block not inoculated. Reprinted from Smith & Read. Mycorrhizal Symbiosis. Plate 2 © (1997) by permission of the publisher Academic Press London. |
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PATH AND SITE OF EXCHANGE |
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Nutrients are obtained by hyphae, which extend from roots, and can therefore penetrate and exploit a larger volume of soil. |
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From uptake by hyphae, nutrients are transferred to the fungal sheath, intercellularly to the hartig net and with the exception of ECM and monotropoid mycorrhizas, intracellularly into plant cells. The fungal sheath can act as a storage site for nutrients, and allowing the fungi to continue to provide nutrients to the plant when soil concentrations decrease. |
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In return for receiving extra nutrients, mycorrhizal plants loose between 10% and 20% of the photosynthates they produce, which go to the formation, maintenance and functioning of the mycorrhizal fungi and their associated structures. (Jakobsen & Rosendahl, 1990). |
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With regard to the transfer of carbon from plant to fungus, labelled carbohydrates entering the fungi appear in the hyphae as trehalose, and in some strains as mannitol. When carbohydrates are transferred from fungus to plant, as in orchid and monotropoid mycorrhizas, the reverse occurs, with trehalose and mannitol being converted to sucrose. |
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In AM mycorrhizas nutrient exchange occurs via the arbuscles. This is an intracellular interface, a situation also found in ericoid mycorrhizas. In contrast, the interface between plant and fungi in ECM takes place via the hartig net, an intercellular structure of varying thickness. |
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| The AM interface (above left) and
the ECM interface (above right), where the exchange of nutrients
occurs. In AM, exchange is via the intracellular arbuscles. In ECM,
it is via the intercellular hartig net. From Allen, M.F. (1992) Mycorrhizal Functioning. An Integrative Plant-Fungal Process. Page 364 figures 11.1A & B. © 1992 Chapman & Hall, London. © 1998 with kind permission of Kluwer Academic Publishers. |
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RATES OF UPTAKE
Direct uptake of nutrients by plant roots can often result in zones of
nutrient deficiency around the root system. Subsequent absorption of
nutrients is then limited entirely by the rate at which nutrients move
through the soil. |
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The rate at which nutrients move through the soil can be crucial to their availability. Phosphorus, in the form of phosphate (PO4-), has very low mobility in soil, and therefore tends to be the limiting nutrient in most ecosystems. In contrast, ammonia (NH4+) is about 10 times more mobile than phosphate, but is also required and absorbed in 10 times greater quantity (Harley, 1991). |
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NITROGEN Nitrate (NO3-) is more mobile than ammonia, and is the main source of nitrogen in most soils. The exception to this rule is in acidic soils, such as those of forests and heathlands, where ammonia is in grater abundance. Most plants will preferentially uptake ammonia over nitrate (Marschner & Dell) and research with ECM fungi has shown that many may not be able to utilise nitrate (Harley, 1991). This is advantageous to plants, as even if ammonia is in low concentration, the ECM fungi will absorb it more rapidly than nitrate (Smith & Read, 1997). Ericoid mycorrhizas are particularly efficient at assimilating ammonia, since their partner plants are mostly heathland plants, which grow on the acidic soils which are high in that form of nitrogen. |
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ECM and ericoid mycorrhizas can also utilise organic N sources, through the production of acid proteinases (Leake and Read, 1990). This allows them access to N sources that are unavailable to non-mycorrhizal plants. In contrast, AM inoculation has little effect on N nutrition, and when nitrate is plentiful in the soil, AM plants have no advantage over non-AM plants (Harley, 1991). In agreement with this, ECM and ericoid plants tend to dominate in N-limited soils, and AM dominate in P-limited soils. |
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PHOSPHORUS Phosphorus (P) is the limiting nutrient in most soils and it can be said that, in general, large growth increases due to mycorrhizal infection are due to increases in P absorption (Bolan et al, 1987). In AM plants, when root growth is restricted, up to 80% of P obtained by the plant can come via the hyphal network (Li et al, 1991). |
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![]() From Kendrick (1999).© Mycologue Publications with permission.. |
Shown left is a classic mycorrhizal experiment showing enhanced growth by the plants on the left which have been inoculated by mycorrhizas, and the control (not inoculated) plants on the right, showing less growth. |
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AM fungi are actually unable to liberate P in this way (Bolan,1991) and as with N in organic complexes, it is ECM and ericoid mycorrhizas that this is particularly well developed in. |
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The beneficial effects of mycorrhizas with regard to P uptake are lost if the concentration of P in the soil increases. Both AM and ECM infection decreases with high P soils. In such soils, the may lead to the mycorrhizal association being abandoned, or can lead to growth depression - essentially a shift from mutualism to parasitism. |
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MONOTROPOID AND ORCHID MYCORRHIZAS The cases of monotropoid and seedling orchid mycorrhizas show different movement of nutrients between plant and fungus. The plants can be considered to be myco-heterotrophic, since they obtain their supplies of organic C from a fungus, but give nothing back in return. Mineral nutrients may also be supplied to the plants by their fungi. However, the fungi appear not to be disadvantaged by this relationship as in monotropoid mycorrhizas, the fungi sequesters its carbon from other photosynthetic plants. Monotropoid mycorrhizas are often ECM with surrounding trees, and carbohydrates form that plant sustain both the fungus and the associated Monotropa sp. The monotropoid relationship has been referred to as epiparasitic, but since the fungi is not damaged in this symbiosis, this term has been dropped in favour of myco-heterotrophic (Smith & Read, 1997). The fungi almost certainly receives nothing in return from Monotropa sp. although whether fungi do in the adult stages of orchid growth remains to be seen. |
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To find out about the other effects of mycorrhizas or the commercial applications, click below: |
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The effects of mycorrhizas |
The commercial applications of mycorrhizas |
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