MYCORRHIZA

The term ‘mycorrhiza’ was suggested by A. B. Frank to characterize the association between fungi and
roots of higher plants. (Myco – fungus and rhiza – root). The structure and composition of mycorrhizae
are variable. Though numerous fungi live in intimate contact with plant roots, only a few enter into
symbiotic association. Mycorrhizal associations are mutualistic: the fungal hyphae help the plant in
absorption of nutrients and water from the soil and intern the plant provides steady supply of carbon and
energy. The mycorrhizal fungal filaments in the soil are truly extensions of root systems (100 to 1000
times) and are more effective in nutrient and water absorption than the roots themselves. Tillage, erosion,
fumigation, etc. eliminates these beneficial fungi.

There are three major types of mycorrhizal associations:

1. ECTOMYCORRHIZA / ECTOTROPHIC MYCORRHIZA

2. ENDOMYCORRHIZA / ENDOTROPHIC MYCORRHIZA
➢ VESICULAR-ARBUSCULAR MYCORRHIZA (aseptate fungi)
➢ ORCHIDACEOUS MYCORRHIZA (septate fungi)
➢ ERICOID / ERICACEOUS MYCORRHIZA (septate fungi)
3. ECTENDOMYCORRHIZA / ECTENDOTROPHIC MYCORRHIZA
ECTOTROPHIC MYCORRHIZA:

• Ectomycorrhizal associations are mostly between:

Host – gymnosperms – pines, oak, firs, beech, sweet chestnut, eucalyptus, etc.
Fungi – basidiomycetes (Agaricus), ascomycetes, phycomycetes
• The fungi associated as ectotrophic mycorrhiza belong to the following families:
Ascomycotina: Tuber, Elaphomyces
Basidiomycotina:
- Hymenomycetes: Amanita, Boletus, Russela
- Gasteromycetes: Rhizopogon, Scleroderma
• The association of the fungus with the roots gives it a stumpy and swollen appearance. The infection
process is stimulated by the production of root exudates which attract the fungi towards the root. The
hyphae aggregate and surround the root externally and this structure is called mantle (or sheath). The
hyphae extending outwards from the mantle help in absorption of nutrients and water.
• The hyphae send branches inwards between cortical cells and extend through the intercellular spaces,
but do not enter the cells. The fungal hyphae that penetrate between the cortical cells is called hartig
net.
• The cortical cells associated with hartig net are larger in size. Hartig net helps to absorb nutrients from
the cortical cells.
• Ectotrophic mycorrhizal fungi grow slowly in a culture media and require additional supply of amino
acids like thiamine and vitamins.
ENDOTROPHIC MYCORRHIZA:

In this case, the fungal hyphae invade the root cortical cells and based on the nature of the fungi, they are
divided into the following groups:

o Aseptate fungi belonging to Zygomycotina:

▪ This type of mycorrhiza is also called Vesicular-Arbuscular Mycorrhiza or Arbuscular
Mycorrhiza (VAM or AM). The VAM fungi are geographically ubiquitous and this type of
association is the most common among all mycorrhizae.
▪ They occur in all groups of plants, that is, Bryophytes, Pteridophytes, Gymnosperms and
Angiosperms (agricultural crops like tomato, cotton, peas, coffee, tea, sugarcane, rubber, coconut,
citrus, shrubs, tropical trees, temperate trees)
▪ VAM fungi mostly belong to the genera – Glomus, Acaulospora, Gigaspora, Entrophospora,
Scutellospora, etc. AM fungi do not produce vesicles, eg. Gigasporacae
▪ The fungi are obligate biotrophs and do not grow on synthetic media.
▪ VAM fungi produce the following characteristic structures:

- EXTERNAL HYPHAE – the external hyphae

are dimorphic – thick walled permanent hyphae and thin
walled short-lived hyphae. The external hyphae bear
various kinds of spores in the soil. These hyphae form
appressoria at the site of penetration into the root.
- CHLAMYDOSPORE – VAM fungi produce
large resting spores that can survive adverse conditions.
- ARBUSCULES – after the infection, the
hyphae form arbuscules in the cortical cells. These are
made of highly branched brush-like structures called
haustoria in cortical cells. These help in the exchange
of nutrients. The cortical cells swell up as a response to
the infection.
- VESICLES – these are spherical or oval thick-
walled structures formed in the intercellular region and
within cortical cells. These are storage organs for fat
granules.
o Septate fungi belonging to Basidiomycotina (rarely Deuteromycotina):
This is further divided into 2 groups

Orchidaceous
Septate fungi - mycorrhizae Arbutoid
Ericoid
Monotropoid

1. ORCHIDACEOUS MYCORRHIZA
- The plants belonging to family Orchidaceae (orchids) are obligately dependent on
mycorrhizal association for germination, as orchid seed has virtually no energy reserve and
obtains its carbon from the fungal symbiont. Growth and cell division is dependent on
external source of carbohydrates supplied by the fungus (Basidiomycetes). Examples of fungi:
Armillaria, Fomes and Rhizoctonia
- The association is said to be TRIPARTITE association → fungi + orchid + endomycorrhizal
host plant
- These fungi are capable of utilizing complex carbohydrates like cellulose and lignin in soil.
- The hyphae do not form a mantle on the exterior surface, but penetrate into the cortical cells
and form coiled structures called arbuscules. These internal hyphae eventually collapse or are
digested.
Coiled hyphae -
Orchidaceous
Mycorrhiza
2. ERICOID MYCORRHIZA
- Ericoid mycorrhizae are formed by Ascomycete fungi and associate with roots of plants
belonging to Ericales (cranberry, blueberry).
- The fungi form hyphal coils within the cortical cells
- The plant’s rootlets are covered with a sparse network of hyphae; the fungus digests
polypeptides saprotrophically and passes absorbed nitrogen to the plant host.
- There are of two types (host plant is the same - Ericales):
a) ARBUTOID
 The host plants - Arctostaphylos and Arbutus
The fungi of arbutoid mycorrhiza are Basidiomycetes. The association is
ectendomycorrhizae
 The fungi form hartig net and penetrate into the cells and form intracellular coils
 Externally they form the mantle or sheath

b) MONOTROPOID
 The fungi like Boletus are associated with achlorophyllous plants like Monotropa
indica (Indian pipe) - ectendomycorrhiza
 The roots are surrounded by dense fungal network called the sheath and extend
outward for absorption of nutrients
 The hartig net surrounds only the epidermal cells and do not enter the cortex region.
However the cortical cell walls invaginate to accommodate the growing hyphae and
form structures called fungal peg (intrusions from the hartig net into the cortical cell
walls are known as fungal pegs (do not enter the cell))

Fungal peg
ECTENDOMYCORRHIZA:

▪ The term ectendomycorrhiza should be used as a purely descriptive name for mycorrhizal roots which
exhibit the characteristics of both ectomycorrhizas and endomycorrhizas. Ectendomycorrhizas are
essentially restricted to the plant genera Pinus (pine), Picea (spruce) and, to a lesser extent,
Larix (larch). (Ectomycorrhiza is converted to endomycorrhiza in older plants → ectendomycorrhiza)
▪ The fungal taxa usually involved are two species of Wilcoxina (W. mikolae, W. rehmii),
Sphaerosporella brunnea.
▪ Ectendomycorrhizas have the same characteristics as ectomycorrhizas but show extensive
intracellular penetration of the fungal hyphae into living cells of the host root.
▪ This association is characterized by a thin mantle (sometimes absent), hartig net, and various degrees
of intracellular hyphal penetration into epidermal and cortical cells soon after hartig net development.
▪ The formation of ectendomycorrhizas begins with formation of a hartig net, which grows behind the
apical meristem of the growing root. Growing up the older parts of the root, intracellular penetration
increases, with the oldest cells being almost filled with coils of septate hyphae, up to 15 mm wide.
▪ The association is stable for at least a year, with no evidence of hyphal degeneration or lysis.
Ectoendomycorrhizal formation induces the growth of short roots, similar to the ectomycorrhizal
association. Emergent roots become covered in a matrix of highly branched hyphae. This coarse
sheath develops behind the apex, between root hairs, and will eventually cover the entire root, except
when the lateral roots grow very rapidly.
▪ The Hartig net penetrates between the epidermal and outer cortical cells and later extends to the inner
cortex. Intracellular penetration occurs a few cells away from those in which the earliest hartig net
formation is seen. Once inside a cell, the hyphae branch repeatedly.
▪ With the exception of the intracellular hyphal penetration, the structure and development of
ectendomycorrhizas are very similar to that seen in the ectomycorrhizal association.
▪ The other examples of ectendomycorrhiza are Arbutoid and Monotropoid types (they form typical
ectomycorrhizas with conifers such as Pinus and Fagus but can also form ectendomycorrhiza with
other hosts).

Hartig net

Mantle/sheath

External hyphae Coiled hyphae

SIGNIFICANCE OF MYCORRHIZA:
❖ Inoculation of mycorrhizal fungi to soils may improve the overall growth of plants.
❖ Infection by mycorrhizal fungi improves nutrient utilization and absorption of Zn, S, and P which
have low mobility in the soil.
❖ Some plants entirely depend on mycorrhizal uptake of phosphorus. Mycorrhizal fungi help in the
uptake of organic and inorganic phosphorus (P) by the production of phytases, phosphatases or
organic acids.
❖ It has been suggested that exudates from mycorrhizal roots may stimulate the atmospheric nitrogen
fixation by free-living rhizosphere microorganisms. The mycorrhizal infection is particularly
important for leguminous trees because of their need for an adequate P supply during nodulation by
Rhizobium.
❖ Both nitrogen fixation in legumes by Rhizobium and enhancement of P uptake by mycorrhizal fungi
can reduce the use of chemical fertilizers (since these chemicals lead to air, water and soil pollution).
❖ The production of binding substances such as exopolysaccharides help in binding the fungal
mycelium together (hold the soil together by forming aggregates) and reduce soil erosion.
❖ Mycorrhizal fungi are not only a major component of soil fertility but also play a significant role in
regulation of soil biological activity.
❖ Mycorrhizal infection reduces susceptibility (or increases tolerance) of plants towards soil-borne
pathogens like Phytophthora, Pythium, Fusarium, etc. and nematodes. The application of
mycorrhizal fungi along with other conventional techniques may be integrated into disease
management in nurseries, plantations, agricultural fields, etc.