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Brian J. Brian C. Philip J. Han Y. Roy V. Professor, University of Northern British Columbia. 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. Endomycorrhizas are variable and have been further classified as arbuscular, ericoid, arbutoid, monotropoid, and orchid mycorrhizas. The fungal hyphae do not in fact penetrate the protoplast i.

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 [5] and DNA sequence analysis [20] suggest that this mutualism appeared million years ago , when the first plants were colonizing land.

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. Ericoid mycorrhizas are the third of the three more ecologically important types.

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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. 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.

All orchids are myco-heterotrophic at some stage during their lifecycle and form orchid mycorrhizas with a range of basidiomycete fungi. This type of mycorrhiza occurs in the subfamily Monotropoideae of the Ericaceae , as well as several genera in the Orchidaceae.

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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. Mycorrhizal fungi form a mutualistic relationship with the roots of most plant species. In such a relationship, both the plants themselves and those parts of the roots that host the fungi, are said to be mycorrhizal.

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The Orchidaceae are notorious as a family in which the absence of the correct mycorrhizae is fatal even to germinating seeds. Recent research into 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 to be hoarding nitrogen from plant roots in times of nitrogen scarcity. Researchers argue that some mycorrhizae distribute nutrients based upon the environment with surrounding plants and other mycorrhizae.

They go on to explain how this updated model could explain 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. The mycorrhizal mutualistic association provides the fungus with relatively constant and direct access to carbohydrates , such as glucose and sucrose. In return, the plant gains the benefits of the mycelium 's higher absorptive capacity for water and mineral nutrients, partly because of the large surface area of fungal hyphae, which are much longer and finer than plant root hairs , and partly because some such fungi can mobilize soil minerals unavailable to the plants' roots.

The effect is thus to improve the plant's mineral absorption capabilities. Unaided plant roots may be unable to take up nutrients that are chemically or physically immobilised ; examples include phosphate ions and micronutrients such as iron.

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One form of such immobilization occurs in soil with high clay content, or soils with a strongly basic pH. The mycelium of the mycorrhizal fungus can, however, access many such nutrient sources, and make them available to the plants they colonize. Another form of immobilisation is when nutrients are locked up in organic matter that is slow to decay, such as wood, and some mycorrhizal fungi act directly as decay organisms, mobilising the nutrients and passing some onto the host plants; for example, in some dystrophic forests, large amounts of phosphate and other nutrients are taken up by mycorrhizal hyphae acting directly on leaf litter, bypassing the need for soil uptake.

In some more complex relationships, mycorrhizal fungi do not just collect immobilised soil nutrients, but connect individual plants together by mycorrhizal networks that transport water, carbon, and other nutrients directly from plant to plant through underground hyphal networks.

Suillus tomentosus , a basidiomycete fungus, produces specialized structures known as tuberculate ectomycorrhizae with its plant host lodgepole pine Pinus contorta var.

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. The mechanisms by which mycorrhizae increase absorption include some that are physical and some that are chemical. Physically, most mycorrhizal mycelia are much smaller in diameter than the smallest root or root hair, and thus can explore soil material that roots and root hairs cannot reach, and provide a larger surface area for absorption.

Mycorrhizas: Anatomy and Cell Biology

Chemically, the cell membrane chemistry of fungi differs from that of plants. For example, they may secrete organic acids that dissolve or chelate many ions, or release them from minerals by ion exchange.

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Mycorrhizal plants are often more resistant to diseases, such as those caused by microbial soil-borne pathogens. These associations have been found to assist in plant defense both above and belowground. Mycorrhizas have been found to excrete enzymes that are toxic to soil borne organisms such as nematodes. When this association is formed a defense response is activated similarly to the response that occurs when the plant is under attack. As a result of this inoculation, defense responses are stronger in plants with mycorrhizal associations.

AMF was also significantly correlated with soil biological fertility variables such as soil fungi and soil bacteria, including soil disease. Although salinity can negatively affect arbuscular mycorrhizal fungi, many reports show improved growth and performance of mycorrhizal plants under salt stress conditions [44]. Recent research has shown that plants connected by mycorrihzal fungi can use these underground connections to produce and receive warning signals. The host plant releases volatile organic compounds VOCs that attract the insect's predators. The plants connected by mycorrhizal fungi are also prompted to produce identical VOCs that protect the uninfected plants from being targeted by the insect.

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. 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. How to cite this article: M. Tahat, Kamaruzaman , Sijam and R.

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Othman, Mycorrhizal Fungi as a Biocontrol Agent. Plant Pathology Journal, 9: DOI: Full Name:. Online First. Current Issue.