Mycorrhiza

http://en.wikipedia.org/wiki/Mycorrhiza#Types_of_mycorrhiza
A mycorrhiza is a symbiotic (generally mutualistic, but occasionally weakly pathogenic) association between
a fungus and the 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.

Types of mycorrhiza[edit]
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.[26][27] Endomycorrhiza includes arbuscular, ericoid, and orchid mycorrhiza, while arbutoid
mycorrhizascan be classified as ectoendomycorrhizas. Monotropoid mycorrhizas form a special category.

Arbuscular mycorrhiza[edit]
Endomycorrhizas are variable and have been further classified as arbuscular, ericoid, arbutoid, monotropoid,
and orchid mycorrhizas.[28]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 balloonlike (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[24] and DNA
sequence analysis[29] 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.[25] The hyphae of arbuscular mycorrhizal fungi produce the glycoproteinglomalin, 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)

Ectomycorrhiza[edit]
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[25] families, orchids,[31] and fungi belonging
to the Basidiomycota, Ascomycota, andZygomycota. 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.[32] An individual tree may have 15 or more different fungal EcM
partners at one time.[33] 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 likely be between 20000 and 25000. [34]
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.[35] 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. [36][37]
The first genomic sequence for a representative of symbiotic fungi, the ectomycorrhizal basidiomycete Laccaria
bicolor, has been published.[38] 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 mycorrhiza[edit]
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.[27]
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.[40]

Arbutoid mycorrhiza[edit]
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. The
difference to ectomycorrhiza is that some hyphae actually penetrate into the root cells, making this type of
mycorrhiza an ectendomycorrhiza

Monotropoid mycorrhiza[edit]
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 nonmutualistic, parasitic type of mycorrhizal symbiosis.

Orchid mycorrhiza[edit]
All orchids are myco-heterotrophic at some stage during their lifecycle and form orchid mycorrhizas with a range
of basidiomycete fungi. Their hyphae penetrate into the root cells and form typical coils

In Agriculture
https://edis.ifas.ufl.edu/pdffiles/AG/AG11600.pdf
a) Increased overall absorption capacity of roots due to morphological and physiological changes in the
plant. There is increased absorption surface area, greater soil area explored (since the fungus acts as an
extension of the root) (Figure 3), greater longevity of absorbing roots, better utilization of low-availability
nutrients, and better retention/storage of soluble nutrients, thus reducing reaction with soil colloids or
leaching losses.
b) Increased mobilization and transfer of nutrients (P, N, S, micronutrients Cu, Zn) from the soil to the
plant. Mycorrhizal fungi have been estimated to substitute up to 500 lb/a of P for citrus and 170 lb/a for
soybeans in tropical areas.
c) Better development of P solubilizing bacteria in the myco-rhizosphere;
d) Increased establishment, nodulation and atmospheric nitrogen fixation capacity in legumes;
e) Modification of plant-pathogen relations: mycorrhizae influence the colonization of roots by other
microorganisms, reduce the susceptibility (or increase the tolerance) of roots to soil-borne pathogens such
as nematodes or phytopathogenic fungi such as Fusarium oxysporum, Fusarium solani, Rhizoctonia solani
and Macrophomina phaseolina. Usually, plants of soybeans, cotton, tomato, oats, and cucumbers are less
susceptible to nematode invasion when they are mycorrhizal. In studies with fungi such as Pythium,
Phytophthora, Fusarium and Verticillium, in most cases (53%) the mycorrhizal interaction is beneficial for
onions, cucumbers, and tomatoes.
f) Secretion of antibiotics and support of a community that competes or antagonizes pathogenic
microorganisms, thus aiding in disease suppression;
g) Increased production of plant growth hormones such as cytokinins and gibberelins;
h) Modification of soil-plant-water relations, promoting better adaptation of plant to adverse environment
conditions (drought, metals). At elevated heavy metal concentrations in soils, mycorrhizal fungi have been
shown to detoxify the environment for plant growth.

In Plant Nutrition
http://www.mycorrhiza.net/benefits.htm
The main benefit is improved uptake of soil phosphorus. Because of better phosphorus nutrition,
mycorrhizal plants can grow much more quickly than non-mycorrhizal plants. The experiments that show
this growth response are done in controlled conditions, and it is unusual for a user in the field to see
responses of the kind that are often shown in scientific (or advertising) photos. The user may see a gain of
a few percent up to double or triple, depending on plant species, soil factors, fertilization, and whether
they already may have native mycorrhizal fungi.
Mycorrhizas appear to have only a minimal effect on uptake of nitrogen, although ECM and ericoids may
have some effect. Mycorrhizas do not fix nitrogen, but some people may confuse this symbiosis with the Nfixing symbiosis between legumes and bacteria of the genus Rhizobium. There are sometimes big effects of
mycorrhizas on certain micronutrients, especially zinc and copper.
Mycorrhizal plants are often more drought tolerant. This is a tricky point, since big mycorrhizal plants in
pots use water much more quickly than little non-mycorrhizal plants in pots. However, it appears that it
can be a very real effect in the ground. This is probably an indirect effect of phosphorus nutrition; plants
fertilized with phosphorus show the same improvement. However, in natural conditions, mycorrhizal
plants are certainly better suited to face dry conditions than non-mycorrhizal plants.
Mycorrhizal plants are more resistant to many root diseases. The scientific results do not all agree, and
tests with certain pathogens have shown the reverse. However, in nature it appears that a range of
beneficial organisms really do fight disease, and those good guys are more abundant if the plants are
mycorrhizal. This explains most of the inconsistent findings, and we can now say with some confidence
that mycorrhizal inoculation is an important part of a holistic disease-fighting program.

In restoration, reclamation, and erosion control, a very important ingredient is the network of mycorrhizal
fungi in the soil. The network builds soil structure, which helps hold the soil together. It also allows survival
of many kinds of seedlings that would otherwise never get big enough to be evident on the job site.
Because some species in the seed mix show up only when inoculated, inoculation in effect increases plant
diversity.
The soil network, in combination with healthy mycorrhizal host plants, is very important in resisting weed
invasion. There have now been a good number of field projects that have successfully fought off weeds,
where other methods that did not involve inoculation have consistently led to nightmare weed
infestations. Inoculation is often not enough in itself, but must work with rapidly growing native plants and
in many cases, some means to temporarily immobilize nitrate, such as a layer of straw or wood chips.

Benefits of Mycorrhizal Biofertilizer

http://www.fnca.mext.go.jp/english/bf/bfm/pdf/4_3_Mycorrhiza0403.pdf
1. Allow plants to take up nutrients in unavailable forms or nutrients that are fixed to the soil. Some plant nutrients,
especially phosphorus, are elements that dissolve were in water in neutral soil. In the extreme acidic or basic soil,
phosphorus is usually bound to iron, aluminum, calcium, or magnesium, leading to water insolubility, which is not
useful for plants. Mycorrhiza plays an important role in phosphorus absorption for plant via cell wall of mycorrhiza to
the cell wall of plant root. In addition, mycorrhiza help to absorb other organic substances that are not fully soluble
for plants to use, and also help to absorb and dissolve other nutrients for plants by storage in the root it is associated
with.
2. Enhance plant growth, improve crop yield, and increase income for the farmers. Arising from improved water and
essential nutrients absorption for plant growth by mycorrhiza, it leads to improvement in plant photosynthesis,
nutrients translocation, and plant metabolism processes. Therefore, the plant has better growth and yield, reduce
the use of chemical fertilizer, sometimes up to half of the suggested amount, which in turn increases income for the
farmers. As in the trial involving mycorrhizal biofertilizer on asparagus it was observed that, when the farmers used
suggested amount of chemical fertilizer together with mycorrhizal biofertilizer, it was found that the crop yield
improved by more than 50%, and the farmers income increased 61% higher than when chemical fertilizer alone was
used.
3. Improve plant resistance to root rot and collar rot diseases. Mycorrhizal association in plant roots will help plant to
resist root rot and collar rot diseases caused by other fungi.
4. It can be used together with other agricultural chemicals. Mycorrhiza are endurable to several chemical
substances; for example; pesticide such as endrin, chlordane, methyl parathion, methomyl carbofuran; herbicide
such as glyphosate, fuazifopbutyl; chemical agents for plant disease elimination such as captan, benomyl, maneb
triforine, mancozed and zineb.

http://www.angelfire.com/hiphop/mycoeducation/mycorrhiza.html
GENERAL BENEFITS OF MYCORRHIZA ASSOCIATION:
1. Increase Drought Resistance; mycorrhizal association increases potential area of water uptake. For
instance, Ectomycorrhiza hyphae can be as much as 1-80m per 1cm of plant root.
2. Increase Resistance to Waterlogging; the external mycelium acts as a protective sheath, and may
even aggregate and differentiate to form structures know as strands, to aid in regulation of water
transfer.
3. Increase Transpiration rate of plant host; increase stomatal conductance and change hormone
balance, as such cells divide more and dont elongate, resulting in a larger biomass with more
branches.
4. Increase Hydraulic Conductivity; results due to extra phosphorous supplied by mycorrhiza
association which is used to help maintain membranes and efficient function and permeability
control.
5. Parasitic Nematode control