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*

Mycorrhizae
Mutualism: Interactions between individuals of different species that
benefit both partners.

* Facultative Mutualism occurs when a species can live without its


mutualistic partner.

* Obligate Mutualism occurs when a species is dependent on a


mutualistic relationship.
Two most common types of mycorrhizae:

Arbuscular mycorrhizal fungi (AMF)


Produces arbuscules - site of exchange
between plants and fungi, hyphae - fungal
filaments, and vesicles - energy storage
organs.

Ectomycorrhizae (ECM)
Forms mantle around roots - important in
increasing plant access to phosphorus and
other immobile nutrients.
Differences
Ectomycorrhizal fungi do not penetrate the cell wall of the plant's
root cells

Arbuscular mycorrhizal fungi penetrate the cell wall formerly called


vesicular-arbuscular mycorrhizae balloon-like (vesicles) or dichotomously
branching invaginations (arbuscules)
Root – fungus association is called Mycorrhiza.

Mycorrhizal plants increase the surface area of the root


system and absorb nutrients from soil especially
phosphorus and micronutrients by huphae that goes
beyond root zone to absorb nutrients.

Almost 90 % of crop plants are mycorrhizal mostly


of AM type.
Therefore, AM association in crop plants plays
significant role in enhancing nutrient mobilization
towards root.
Mycorrhiza increase the surface area of the root
Absorb nutrients from soil especially phosphorus
and micronutrients through hyphae and mobilize
into the host cell.
Type of mycorrhizae
Ectomycorrhizae (EM)
• The fungal mycelia extend inward, between root cortical cells, to form a
network (Hartig net) and outward into the surrounding soil. Usually the
fungal hyphae also form a mantle on the surface of the root.

• Hyphae do not penetrate into cells but contact with roots is very close
and metabolites are transferred in both direcions.

• Some ectomycorrhizae produce large above ground sporocarps or


mushrooms which facilitate dispersal of spores along with underground
fruiting bodies.

• Found on many woody plants ranging from shrubs to forest trees. Host
plants belong to the families Pinaceae, Fagaceae, Betulaceae and
Myrtaceae and a few others but no grasses.
*
*fungal component are mainly
basidiomycetes, but some
ascomycetes

*at least 5,000 species of fungi


involved in ectomycorrhizal
relationship2
Ectomycorrhizae predominate in temperate and boreal forests

Organic soils
The fungal hyphae form a tissue-like sheath (mantle) on the surface
of the nutrient- absorbing rootlets.

Ectomycorrhizae have been shown to degrade proteins in soil organic


matter, obtaining a source of nitrogen that would otherwise be
unavailable to plants.
*
Hyphae extend into the soil, and also grow between the root cortical cells to
form a 'Hartig net‘.

Mantel

Hartig Net
* the function of the
ectomycorrhizal
mutualism depends on
the ability of the fungal
symbionts to take up
nutrients

* the symbiotic nutrient


exchange plays a major
role in plant nutrition as
well as in resistance of
plants against pathogens,
heavy metals, drought
stress, etc.3
www.mycorrhizae.com/images
*
* Regulated gene expression is
an important mechanism for
controlling ectomycorrhizal
symbiosis development and
functioning.

* Gene profiling studies have


identified genes that are
required for fungal
attachment, plant defense,
and symbiosis-related
metabolism.4
Type of mycorrhizae
Endomycorrhizae
• Mycorrhizal association with intracellular penetration of the host root
cortical cells by the fungus as well as outward extension into the
surrounding soil.

• The fungus initially grows between cortical cells, but soon penetrates
the host cell wall and grows within the cell.

• Common in must species of herbaceous angiosperms, flowering plants,


annual and perennial crops, and many of the gymnosperm genera.
Type of mycorrhizae
Endomycorrhizae
• Consists primarily of vesicular-arbscular mycorrhizae (VAM),
especially common in grasslands.

• Vesicles are rounded intercellular


structures which act as storage
organs

• Arbuscles are branched intercellular


structures, which serve as the site of nutrient transfer between the
fungus and the root.
*found in the roots of
roughly 70% of all plant
species
*fungal component are
Glomales
*hyphae penetrates the
outer cells of plant
root1
Arbuscular mycorrhizae is the dominant type in the
tropics, and in grasslands and deserts of temperate
latitudes

Mineral soils
Arbuscular mycorrhizae are formed only by fungi in the
division Glomeromycota

Typically associated with the roots of herbaceous


plants, but may also be associated with woody plants
Hyphae originate from the colonized host plant root

Increase water and nutrient uptake

Small enough to invade smaller pores than roots

Some hyphae invade neighboring plants
Lecture
Spores
• Swollen structures with one or more subtending hyphae that form in the soil or
in roots. Spores usually develop thick walls, which often have more than one
layer. They can function as propagules. Spores of VAM fungi are sometimes
called chlamydospores or azygospores.

• The tiny spores of EM fungi are usually borne in above ground (i.e.
mushrooms) or below ground (i.e. truffles) reproductive structures. They are
dispersed by wind, water, and animals that may eat the fungal fruiting body.

• The much larger spores of VAM fungi are often borne directly in the soil,
formed inside the root, or clustered in various ways with other spores.VAM
spores are less likely than EM spores to be dispersed by wind.

• More often, they are moved with soil or sediment in water, or carried by
earthworms and other vectors.
Hyphae
• hyphae: thread or filament forming the mycelium of fungus.

• The hyphae of mycorrhizal fungi can make most of the soil's microbial
biomass.

• The hyphae of VAM fungi are usually recognizably distinct from other
kinds of soil fungi. These hyphae are relatively large, with smaller side
branches that arise from characteristic bumps and angles on the main trunk
hypha. The hyphae have few cross-walls (septa) and the larger ones persist
in the soil long after they are dead.

• Hyphae of EM fungi and some others may aggregate to form coarse hyphal
strands (rhizomorphs). These can carry water and nutrients a considerable
distance toward the plant.
Mycorrhizae inoculum
• Mycorrhizal inoculum is material that carries mycorrhizal fungi in a
usable form to the intended host plants.

• In the case of VAM fungi, inoculum may be spores, hyphae, and


colonized root fragments.

• The quality of mycorrhizal inoculum is not easy to assess, but the best
way is an expression of the number of propagules per unit volume of
material. Propagules include spores, hyphae, and root fragments.
Functions of Mycorrhizal
• Fungus uses organic nutrients (including sugars) produced by plant
ECM are not obligate mutualists but VAM are.

• Fungi impart benefits to plant by increasing: nutrient absorption,


especially phosphorus, water uptake, tolerance of harsh
environmental conditions, including polluted environments
(e.g. acts as shield against acidity, elemental toxicity and pathogens).
Factors that can affect of
mycorrhizal formation
• > pH of soil----------------------------------------(-)
• > soil moisture------------------------------------(-)
• > soil depth----------------------------------------(-)
• > amount of above ground plant cover--------(+)
• > grazing-------------------------------------------(-)
• > pesticides----------------------------------------(-)
• > soil fertility--------------------------------------(-)
Benefits of Mycorrhizae
• Increased uptake of nutrients
– Hyphae explore the soil for nutrients, increase surface area for nutrient
absorption transport them back to the plant.
– The nutrients P, Zn, C, N, Cu and S have been shown to be
absorbed and translocated to the host by mycorrhizal fungi

• Increased rootlet size and longevity


– Mycorrhizal plants have larger roots than nonmycorrhizal plants
regardless of whether mycorrhizal fungi are present.
Benefits of Mycorrhizae
• Water relations
– Hyphae explore the soil for water and increase surface area for
absorption
– Some mycorrhizae alter the plant’s physiology, increasing stomatal
resistance, resulting in less water loss.

• Improved growth rate


Benefits of Mycorrhizae
• Tolerant of harsh conditions

– fungi are more tolerant of acidity, elemental toxicity and high soil
temperatures than are higher plants and able to, in some cases
(ectomycorrhizae), shield the root from these condition.

– Lower levels of heavy metals generally found in mycorrhizal


plants than nonmycorrhizal plants.
Lecture
Benefits of Mycorrhizae
• Increased seedling survival
– Mycorrhiza promotes plant survival, whether new seedlings or out-
planted container stock.

– Survival of inoculated plants can be up to five times the survival of


uninoculated plants.

– Improved survival is no doubt due to a combination of mycorrhizal


benefits, including faster growth to help overtop weeds, protection
from pathogens, and improved drought tolerance.
Mycorrhizae and Plant diversity
• Biodiversity of belowground fungal symbionts increases biodiversity of
above ground plants

• Increased access to nutrients becomes restricted under competitive conditions

• Differences in functional capacity of a specific fungus-plant combination


appear to explain the effect

• In ecosystems, increased functional capacity allows one plant species to


perform better than others

• Restored plant communities have been found to be more diverse when


mycorrhizal fungi are present when both inoculated and uninoculated areas
receive the same seed mix.
Mycorrhizae and Plant diversity
Basis for fungal species richness on plant biodiversity and production

No symbionts
Mycorrhizae and Plant diversity
Basis for fungal species richness on plant biodiversity and production

No symbionts One symbiont

Increasing diversity
Mycorrhizae and Plant diversity
Basis for fungal species richness on plant biodiversity and production

No symbionts One symbiont Two symbionts

Increasing diversity
Increasing productivity
Mycorrhizae and Plant diversity
Basis for fungal species richness on plant biodiversity and production

No symbionts One symbiont Two symbionts Four symbionts

Increasing diversity
Increasing productivity
Lecture
Mycorrhizae and pathogens
• Mycorrhizal fungi have long been known to help defend roots against soil-
borne disease.

• Mycorrhizae promote beneficial bacteria that may be directly responsible for


protection against root pathogens.

• Mycorrhizal fungi may reduce the incidence and severity of root diseases.
The mechanisms proposed to explain this protective effect include:
– development of a mechanical barrier-especially the mantle of the EM-to infection
by pathogens
– production of antibiotic compounds that suppress the pathogen,
– competition for nutrients with the pathogen, including production of siderophores
– induction of generalized host defense mechanisms.
Disadvantage of Mycorrhizae

• When the nutrient levels in the soil are so low and the fungus cannot
extract extra nutrients and no benefit accrues to the plant in return for
carbon transferred to fungus parasitism may be occur.

• Sometimes plant growth suppression has been attributed to


mycorrhizal colonization, but usually this occurs only under unusual
circumstances that affect plant photosynthesis (low light and cold
temperatures) or high-phosphorus
Mycorrhizae and Succession
• Many below-ground processes, including plant-microbial interactions
influence the rate and direction of succession, whether primary or
secondary.

• Formation of mycorrhizae as one of the ten major successional


processes that determines change in species composition during
succession.

• Mycorrhizal fungi alter succession and community composition by


altering the relative success of plants, both in the seedling stage and
during later development
Mycorrhizae and Succession
• Succession on arid and semiarid lands is influenced in part by the
relative density of mycorrhizal inoculum.

• The nonmycorrhizal of the early stages of succession are initially


replaced by facultative mycotrophs and finally by obligate mycotrophs.
Mycorrhizae less common in the early seral stages
More prevalent in low fertile sites and harsher environments
Many climax species are dependent on mycorrhizae
Mycorrhizae and Soil
Structure
• Soil structure is tremendously important in the health of
the vegetation, because it facilitates water infiltration,
aeration, root growth, and movement of soil animals.

• Structure refers to the arrangement of soil particles into


aggregates. Mycorrhizal fungi are key players in soil structure
• by their hyphae. Pore space allows ready movement of water, air, roots,
and animals. Soils without pore space are very unfriendly to plant growth.

• Both ECM and VAM fungi facilitate soil structure by binding soil
aggregates together, to each other, and to plant roots

• Soil bacteria cement soil particles and contribute importantly to soil


structure, and mycorrhizal hyphae help supply some of the fuel that keeps
the soil bacteria alive
Mycorrhizae and Weeds
• Healthy native ecosystems are resistant to weed "ruderal" invasion; an
effect promoted by vigorous, mycorrhizal native plants.

• Many weeds do not need to become mycorrhizal, and can grow in places
where natives cannot become mycorrhizal. In such locations, weeds win
the competitive struggle greatly incapacitated natives.

• Weeds usually require relatively higher soil fertility than native species.
Healthy, mycorrhizal natives rapidly remove mobile nutrient ions from the
soil, making the site inhospitable to weeds.

• Some weeds react to mycorrhizal fungi as they would to root pathogens.


Seedlings of Russian thistle, an important weed in the western US, soon
die if the soil is full of mycorrhizal hyphae. Other weed species that are
intolerant of mycorrhizal fungi may be discovered in the future.
Management of Mycorrhizae

• Most of the mycorrhizal fungi are in the top 15 cm of the soil


• Preserve topsoil and microbial activity
• Do not use chemicals that kill fungi
• Do not over-fertilize
• Keep native plants
The End

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