Professional Documents
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Leccinum aurantiacum, an
ectomycorrhizal fungus
Wheat is in a arbuscular
mycorrhiza relationship
Mycorrhiza
2. Ectomycorrhizal fungi
• Fungus remains on the outside of plant cells (Parniske,
2008)
• Mainly found with most of the economically important
tree species in temperate regions, and in some tropical
tree species Ectomycorrhiza (left)
Endomycorrhiza (right)
• Eg. of plant species: Birch, Eucalyptus, Oak, Pin
• Eg. of fungal families: Basidiomycota, Ascomycota,
Zygomycota
Ectomycorrhiza – Ecology & Distribution
• Ectomycorrhiza are confined to a limited number of
vascular plant genera
• Mostly gymnosperms and a few angiosperms
• Especially in temperate regions
• Many fungi involved in these associations are
cuturable and hence more amenable to experimental
manipulation than VAM fungi
Features
• In contrast to AM, the fungi forming ectomycorrhiza Extraradical mycelia (white) on the roots of
come from a wide spectrum of taxonomic groups Picea glauca (brown)
• Additionally, the range of potential host plants is
smaller [total no. of fungal species (5000) outnumber
woody hosts (2000)]
• Largest number of species belong to Basidiomycotina
(majority of species form such associations with higher
plants)
• In the Ascomycotina, many of the ectomycorrhizal
symbiots form characteristic hypogeous fruit bodies, or
truffles Ectomycorrhizal symbiosis,
showing root tips with fungal
mycelium from the genus Amanita
Orders of Fungi Involved in Ectomycorrhizal
Associations
1. Zygomycotina
2. Endogonales
3. Ascomycotina
4. Elaphomycetales
5. Pezizales
6. Tuberales
7. Basidiomycotina
8. Agaricales
9. Aphyllophorales
10. Gautieriales
11. Hymenogastrales
12. Lycoperdales
13. Melanogastrales
14. Phallales
15. Russulales
16. Sclerodermatales
Ectomycorrhiza – Morphology
• 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
• 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
• When compared to non-mycorrhizal fine roots, ectomycorrhiza may contain very high
concentrations of trace elements, including toxic metals (cadmium, silver) or chlorine
AM/VAM – Ecology & Distribution
• AM are the most common of all mycorrhiza
• Involvement in as many as 90% of all vascular plants
• However, only a few genera and species of fungi form
this association
• These fungi are obligate biotrophs, hence establishing
in vitro cultures is almost impossible
• With a few exceptions, almost all genera of AM fungi
produce vesicles
• Studies and observations have confirmed that members
Vesicular Arbuscular Mycorrhiza
of the genera Scutellospora and Gigaspora form (VAM)
mycorrhizas with arbuscules, but not with vesicles
• VAM association is found in a wide range of habitats,
i.e. from the arctic region to tropical regions
Six genera of fungi in Endogonaceae
• Fungal hyphal networks can contain over 100 meters which have been shown to form
of hyphae in 1 cubic centimeter of soil mycorrhizal associations: Glomus,
Gigaspora, Acaulospora,
Entrophospora Sclerocystis and
Scutellospora
AM/VAM – Spore morphology
AM – With and without colonization
Features
• Stable compound
• Insoluble in water
• Resistant to degradation by heat
• (albeit degrade at very high temperatures; i.e. around 120 oC)
• May last for more than 40 years
• Produced in large amounts
• Resistant to decay
Functions:
• Glues together soil aggregates
• Soil carbon and/or nitrogen storage Fig: Microscopic view of AM growing on a
• Increases water retention near roots corn root. Round bodes are spores. Thread
• Improves nutrients cycling like filaments are hyphae; Coating on the
hyphae is glomalin (bright green spots)
• Improves root penetration by reducing compaction
• Protect hyphae from nutrient loss
Glomalin Production – “Soil binding glue”
MYCORRHIZAL FUNGI
• Up to 20% of a plant’s fixed C is transferred to the mycorrhizal fungus in exchange for water
and nutrients such as N and P (Hodge and Storer, 2015)
• Phosphorous movement in mycorrhizal plants has been found to be 3 to 5 times faster than
their non-mycorrhizal counterpart
• Research have also shown that AM fungi also play major roles in N uptake for their host plants
• The uptake of inorganic and organic forms of N by the fungi continue to benefit the host plant
(Hodge and Storer, 2015)
• Mycorrhizal plants show better responses and tolerance towards drought stresses
Nutrient Uptake & Transport
• Hyphal networks are very efficient at exploring the soil and bringing in phosphate,
Zn, and ammonium (Gerlach et al., 2015)
• Phosphorous is perhaps the most important nutrient made available by mycorrhizal
associations
• Hyphal networks can help extend the plant’s availability to bring in more P from the
solution P pool in the soil (Gerlach et al., 2015)
• They are also able to explore small pores in the soil not accessible by plant roots
(Jakobsen et al., 2005)
Nutrient Uptake & Transport
Nutrients taken in by plants that are infected and by those that are
not infected with AM fungi when no P is added to corn
Corn – Impacts of AM fungi colonization
It is unlikely that mycorrhizal fungi have an ability to directly liberate the phosphorus in
these insoluble inorganic or organic sources, but their ability to infiltrate the soil with
their mycelium means that a greater proportion of the soluble phosphorus is absorbed,
leading to a shift in the equilibrium towards the release of bound phosphorus from the
soil reserves
Mycorrhiza - benefits to plants (Research
Observations)
Mycorrhiza - benefits to plants (Research
Observations)
Effects of mycorrhizal colonisation and P nutrition on growth and P concentrations of roots and shoots
of Trifolium subterraneum (31 to 35 days old)
Mycorrhiza - benefits to plants (Research
Observations)
(a) Dry weight changes and (b) P contents of Allium cepa, colonized by four VA mycorrhizal fungi,
compared with controls (C). YV, Glomus mosseae; LAM, G. macrocarpus var. geosporus; MIC, G.
microcarpus; BR, Scutellospora (Gigaspora) calospora.
Mycorrhiza – Improving crop productivity
• Enhance productivity and quality of crops
• For example, AMF-colonized strawberry exhibited increased levels of secondary metabolites
resulting in improved antioxidant property (Castellanos-Morales et al., 2010)
• AMF can enhance the dietary quality of crops by affecting and production of carotenoids and
certain volatile compounds (Hart et al., 2015)
• Bona et al. (2017) observed beneficial effects of AMF on the quality of tomatoes
• In another study, Zeng et al. (2014) have reported increased contents of sugars, organic acids,
vitamin C, flavonoids, and minerals due to Glomus versiforme resulting in enhanced citrus
fruit quality
• Mycorrhizal symbiosis induces enhanced accumulation of anthocyanins, chlorophyll,
carotenoids, total soluble phenolics, tocopherols, and various mineral nutrients (Baslam et al.,
2011)
• AMF have been employed in a large-scale field production of maize (Sabia et al., 2015), yam
(Lu et al., 2015), and potato (Hijri, 2016), confirming that AMF possess a considerable potential
for enhancing crop yield
Mycorrhiza – Role in Plant defense (Biocontrol)
• Both ectomycorrhiza and VAM have been implicated as agents of biological control through the
inhibition of infection of host plants by certain root-infecting pathogens.
• Like most examples of biological control, however, mycorrhiza never confer complete
immunity against any disease or pest due to the imprecision of biological agents
• There is often a problem again in deciding whether disease resistance is mediated by the
mycobionts directly or is a consequence of the improved nutrient status of the mycorrhizal
plants
• Protective effects of some ectomycorrhiza has been well established and their roles might be
easier to understand than that of VAM; Ex: Protection against Phytophthora cinnamomi in
Pinus patula due to its ectomycorrhizal association (protection achieved through hyphal
sheaths; as no evidence of antibiosis is demonstrated and occurrence of infection when sheaths
are partially formed or removed)
Possible mechanisms of protection:
1. Secretion of antibiotics that inhibit pathogen growth
2. Hyphal sheath act as a physical barrier to penetration
3. Surplus nutrients in and around roots are utilized, thus reducing the amounts available for other
pathogens
4. Induction of host defense mechanisms
5. Sheath supports a protective microbial rhizospher population
Mycorrhiza – Impacts on soil
• Increase soil aggregation and persistence
• Improve water percolation and retention
• Improve nutrient access and mobility
• Increase soil organic matter content
• Reduce soil erosion and improve soil aeration
• Improve plant nutrient status reduces fertilizer input and cost
• Trigger plant defensive systems for disease, pest and abiotic stresses: reduce pesticide need and
cost
• Help alleviate metal toxicity in soil
• Reduce drought and salinity stress
• Extend and expand plant roots functional setup
Effect of Agricultural Practices on Mycorrhizal
Populations
Agricultural practices influence changes in soil which directly affects the status of
mycorrhizal fungi present there
Mentioned below are some of the management practices in agriculture that have
significant impacts on mycorrhizal fungal population
1. Fertilization
2. Tillage
3. Crop Rotation
Effect of Agricultural Practices on Mycorrhizal
Populations
1. Fertilization
• Repeated phosphorous application (P) can inhibit AM fungal development
• Increasing availability of nutrients tend to influence plants to rely less on AM fungi
• With decreasing dependencies, the diversity and population of AM fungi tend to
decrease
• Increased fertilization influence some AM fungal species more than others
• This is demonstrated by a shift in AM fungi species under different crop management
practices
Effect of Agricultural Practices on Mycorrhizal
Populations
2. Tillage
• Soil tillage practices generally destroy the extraradical
mycelial networks of existing AM fungi
• Tillage practices also influence a shift in AM fungal
species population
• Example: large amount of Glomeraceae species found
in tilled soil around the world
• This is also supported by the fact that gloamlin levels
increase in soil after adopting no-till practices
• Research from Maryland Agricultural Research Center
showed an increase inglomalin levels after the start of
no-till
• Glomalin levels were 1.3 milligrams per gram (mg/g)
of soil in year one and after three years of no-till, they
were 1.7 mg/g
• In an adjacent field that was plowed each year, the
glomalin levels were 0.7 mg/g and in a nearby 15-year
grass buffer strip the glomalin level was 2.7 mg/g
Effect of Agricultural Practices on Mycorrhizal
Populations
3. Crop Rotation
• Monocultures can be very harmful to AM fungal
populations and diversity
• This can be especially true when crops are not
highly dependent on the mycorrhizal symbiosis
(e.g. wheat), or crops that are nonmycorrhizal
hosts (e.g. canola) are grown in succession
• Less diversity in crop rotation can affect the AM
fungal community to be both less diverse and less
parasitic, meaning they are less aggressive at
colonizing the host plant
• The impact of crop rotation is crucial to the
diversity and population of AM fungi
Mycorrhizal Biofertilizers gaining traction