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Mycorrhizae
Plant roots and fungi
Mycorrhizae
• Widespread interactions between fungi
and plant (primarily vascular plants) roots
• For angiosperms, gymnosperms, ferns
and some mosses – mycorrhizal
association appears to be the norm
• Range over broad spectrum of
interactions
– Fungus parasitizes plant
– Plant parasitizes fungus
– Most cases – mutualistic – both benefit
Types of mycorrhizae
• Ectomycorrhizae (ectotrophic, sheathing)
– hyphae of fungus do not penetrate cells
of plant root
• Endomycorrhizae – hyphae penetrate
cells of plant
– Arbuscular mycorrhizae (AM) – aseptate
hyphae, most widespread
– Septate hyphae
• Ericoid, Arbutoid & Monotropoid – plants are
Ericales
• Orchid – plants are orchids
Ectomycorrhizae
• Most conspicuous and easily recognized
• Best characterized
• Plant roots are enclosed by a sheath of
fungal hyphae – fungal mycelium penetrates
between cells in cortex of the root
• Fungal tissue may account for up to 40%
mass of root
• Hyphae also extend out into the soil –
extramatrical hyphae
Ectomycorrhizae
• Contains a fungal
sheath
• Parenchyma of root
cortex is surrounded
by hyphae – Hartig
net
Ectomycorrhizal root
Ectomycorrhizae
• Absorbing roots are those that are affected
• Become thicker and repeatedly branched after
infection
Ectomycorrhizae
Ectomycorrhizae Symbionts
• 2000 plant species – primarily temperate
trees and eucalyptus
• Major species of coniferous and
deciduous trees
• Rare to find uninfected trees
• In some trees, the association is obligate,
in others facultative
• Mycorrhizal association important in
forestry
Ectomycorrhizae Symbionts
• Basidiomycetes – Agaricales (many
mushroom species), Lycoperdales,
Sclerodermatales, few Aphyllophorales
– Pisolithus tinctorus – used to form commercial
inoculum for nursery trees, common in
southern pine
• Ascomycota – Pezizales – cup fungi and
truffles
• Over 5000 species of fungi have been
shown to form ectomycorrhizae
Specificity of association
• Great deal of variability
• Most tree species form mycorrhizal
associations with a number of different fungal
species
• May have different mycorrhizal fungi on roots
of one plant
• Some fungi are fairly specific and will form
associations with only one plant species –
these mushrooms are common in stands of
that tree
• Others are not specific
Specificity
• Douglas fir has been
extensively studied
and ca 2000
species of fungi
have been identified
from its roots
• In forests, a high
percentage of
fruiting bodies are
mycorrhizal fungi
Occurence
Methods for detection
• Census of fruiting bodies produced by
different species
• Soil cores – separate and identify
mycorrhizal roots by morphology, Hartig
net
• Recently molecular methods have been
used to identify the fungi present in
mycorrhizal roots – e.g. RFLP
Ectomycorrhizal fungi
• Can also grow saprotrophically
• Many have been cultured
• Most that have been studied do not have
the capability to degrade complex plant
polymers (e.g. cellulose and lignin)
• Depend on soluble carbohydrates
• Many have organic growth factor
requirements – vitamins, amino acids
• Not decomposers but depend on plant
Benefits to fungus
• Provided with source of C and energy
• Plants provided with 14
CO2 demonstrated
that 14
C appears in fungus
• Sucrose from plant converted into trehalose,
mannitol by fungus
• Estimates that up to 10% (or more) of
photosynthate produced by trees is passed
to mycorrhizae and other rhizosphere
organisms
Benefits to trees
• Numerous studies
have shown that tree
growth is better when
mycorrhizae are
present
Benefits to trees
Benefits to trees
• Fungi increase supply of inorganic
nutrients to tree
• P is insoluble in most soils
• Extramatrical hyphae extend over a larger
volume of soil than roots can – increase
ability to absorb insoluble nutrients such
as P
Extramatrical
hyphae
Volume of soil explored
Benefits to trees
• Plant hormones produced by fungus
changes the physiological state of roots –
physiologically active root area for nutrient
and water absorption is increased
• Increases tolerance of plant to drought,
high temperatures, pH extremes, heavy
metals
• Increases resistance to infection by root
pathogens – provides a physical barrier
Ectomycorrhizae
• Mutualistic symbioses – both organisms
benefit from association
• Currently, seedlings in nurseries
inoculated with fungi so that when planted,
they will have better chance of success
Arbuscular mycorrhizae
• AM – much less known about these
associations than about ectomycorrhizae
• Appear to be the most common type of
mycorrhizal association with respect to the
number of plant species that form them
• Found in species in all divisions of
terrestrial plants – widely distributed in
annuals, perennials, temperate and
tropical trees, crop and wild plants
• Estimated to occur on 300,000 plant spp.
Arbuscular mycorrhizal fungi
• All are in the Zygomycota in the Glomales
– or newly proposed phylum
Glomeromycota
• Include ca 130 species in 6 genera
• All are obligate biotrophs
• Form large spores that superficially
resemble zygospores, but not formed from
fusion of gametangia – azygospores or
chlamydospores
• Spore diameters range from 50 to 400 μm
Spores
Arbuscular mycorrhizae
Fossils of spores
found that are as
old as first land
plants – 460 mya
Specificity
• Few species of fungi and many species of
plants – very low specificity
• One fungal species may form association
with many different plant species
• Much different than biotrophic parasites
that have a limited host range
Morphology
• Root morphology is not modified
• To detect, must clear and stain root to
observe fungal structures
• Fungi form both intercellular and
intracellular hyphae
• Intracellular hyphae analogous to
haustoria – called arbuscules – tree like
branching pattern
• Thought to be site of nutrient exchange
between fungus and plant
Arbuscules
• Surrounded by
plant cell
membrane
• Typically
disintegrate after
ca 2 weeks in plant
cell and release
nutrients
• Thought to be site
of nutrient
exchange
Vesicles
• Intercellular hyphae may also form large
swellings – vesicles – at ends of hyphae or
intercalary
• Typically rich in lipids & thought to be involved in
storage
AM
Arbuscular mycorrhizae
• Not as well characterized as
ectomycorrhizae
• Root is not altered in morphology – difficult
to determine when roots are infected –
must clear and stain followed by
microscopic examination
• Fungi are obligate biotrophs – cannot be
grown in axenic culture – so difficult to
conduct experiments
Interaction
• Fungus receives organic nutrition from plant –
since they are biotrophs, don’t know what their
requirements are
• Fungus produces extramatrical hyphae that take
up inorganic nutrients from soil – particularly P,
may also supply N as they may produce
proteinases
• Increase drought tolerance – many common
desert plants are heavily mycorrhizal
• May also increase resistance to root pathogens
Effect of AM
• Growth of plants that are infected better –
particularly if soil is poor in nutrients
Other types of mycorrhizae
• Orchids – orchid seeds are very small and
do not contain enough organic reserves to
allow development of the plant
• Must be infected soon after germination –
fungus provides seedling with carbohydrates
• Basidiomycetes involved in this mycorrhiza
are litter decomposing species of
Rhizoctonia, Armillaria that produce
cellulases
Orchid mycorrhizae
• Fungi are widely distributed outside the
symbiosis – some are plant pathogens,
others are saprotrophs
• Appears to be a delicate balance between
plant and fungus
• Orchid keeps fungus in check by digesting
intracellular hyphal coils, production of
antifungal substances so fungus doesn’t
kill the orchid
Orchid mycorrhizae
• Not clear about
benefits to fungus –
may obtain amino
acids and vitamins
from orchid
Ericoid mycorrhizae
• Plants are Ericaceae – Erica,
Vaccinium - heathland plants
• Fungi are Ascomycota and
Deuteromycota
• Form loose network on
surface & hyphal coils inside
epidermal cells of hair roots
where nutrient exchange is
thought to take place
• Shown to supply N to plant –
fungi secrete proteinases
Arbutoid mycorrhizae
• Plant are also Ericaceae – Arbutus,
Arctostaphylose, Pyrola
• Fungi are basidiomycetes that also form
ectomycorrhizae
• Fungi form sheath and Hartig net, hyphae
also penetrate outer coritcal cells
Monotropoid mycorrhizae
• Plants are nonchlorophyllous
– Monotropa
• Fungi are basidiomycetes –
boletes that form
ectomycorrhizae with other
plants (conifers)
• Plant depends on its
mycorrhizal fungus - for its
organic nutrients as well as
inorganic nutrients
Mycorrhizae
• Key components of ecosystems
• Link plants within a habitat
• Labelled CO2 fed to tree can be found in
seedlings growing nearby
• Retain and conserve mineral nutrients

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Mycorrhizae SLIDESHARE

  • 2. Mycorrhizae • Widespread interactions between fungi and plant (primarily vascular plants) roots • For angiosperms, gymnosperms, ferns and some mosses – mycorrhizal association appears to be the norm • Range over broad spectrum of interactions – Fungus parasitizes plant – Plant parasitizes fungus – Most cases – mutualistic – both benefit
  • 3. Types of mycorrhizae • Ectomycorrhizae (ectotrophic, sheathing) – hyphae of fungus do not penetrate cells of plant root • Endomycorrhizae – hyphae penetrate cells of plant – Arbuscular mycorrhizae (AM) – aseptate hyphae, most widespread – Septate hyphae • Ericoid, Arbutoid & Monotropoid – plants are Ericales • Orchid – plants are orchids
  • 4. Ectomycorrhizae • Most conspicuous and easily recognized • Best characterized • Plant roots are enclosed by a sheath of fungal hyphae – fungal mycelium penetrates between cells in cortex of the root • Fungal tissue may account for up to 40% mass of root • Hyphae also extend out into the soil – extramatrical hyphae
  • 5. Ectomycorrhizae • Contains a fungal sheath • Parenchyma of root cortex is surrounded by hyphae – Hartig net
  • 7. Ectomycorrhizae • Absorbing roots are those that are affected • Become thicker and repeatedly branched after infection
  • 9. Ectomycorrhizae Symbionts • 2000 plant species – primarily temperate trees and eucalyptus • Major species of coniferous and deciduous trees • Rare to find uninfected trees • In some trees, the association is obligate, in others facultative • Mycorrhizal association important in forestry
  • 10. Ectomycorrhizae Symbionts • Basidiomycetes – Agaricales (many mushroom species), Lycoperdales, Sclerodermatales, few Aphyllophorales – Pisolithus tinctorus – used to form commercial inoculum for nursery trees, common in southern pine • Ascomycota – Pezizales – cup fungi and truffles • Over 5000 species of fungi have been shown to form ectomycorrhizae
  • 11. Specificity of association • Great deal of variability • Most tree species form mycorrhizal associations with a number of different fungal species • May have different mycorrhizal fungi on roots of one plant • Some fungi are fairly specific and will form associations with only one plant species – these mushrooms are common in stands of that tree • Others are not specific
  • 12. Specificity • Douglas fir has been extensively studied and ca 2000 species of fungi have been identified from its roots • In forests, a high percentage of fruiting bodies are mycorrhizal fungi
  • 14. Methods for detection • Census of fruiting bodies produced by different species • Soil cores – separate and identify mycorrhizal roots by morphology, Hartig net • Recently molecular methods have been used to identify the fungi present in mycorrhizal roots – e.g. RFLP
  • 15. Ectomycorrhizal fungi • Can also grow saprotrophically • Many have been cultured • Most that have been studied do not have the capability to degrade complex plant polymers (e.g. cellulose and lignin) • Depend on soluble carbohydrates • Many have organic growth factor requirements – vitamins, amino acids • Not decomposers but depend on plant
  • 16. Benefits to fungus • Provided with source of C and energy • Plants provided with 14 CO2 demonstrated that 14 C appears in fungus • Sucrose from plant converted into trehalose, mannitol by fungus • Estimates that up to 10% (or more) of photosynthate produced by trees is passed to mycorrhizae and other rhizosphere organisms
  • 17. Benefits to trees • Numerous studies have shown that tree growth is better when mycorrhizae are present
  • 19. Benefits to trees • Fungi increase supply of inorganic nutrients to tree • P is insoluble in most soils • Extramatrical hyphae extend over a larger volume of soil than roots can – increase ability to absorb insoluble nutrients such as P
  • 21. Volume of soil explored
  • 22. Benefits to trees • Plant hormones produced by fungus changes the physiological state of roots – physiologically active root area for nutrient and water absorption is increased • Increases tolerance of plant to drought, high temperatures, pH extremes, heavy metals • Increases resistance to infection by root pathogens – provides a physical barrier
  • 23. Ectomycorrhizae • Mutualistic symbioses – both organisms benefit from association • Currently, seedlings in nurseries inoculated with fungi so that when planted, they will have better chance of success
  • 24. Arbuscular mycorrhizae • AM – much less known about these associations than about ectomycorrhizae • Appear to be the most common type of mycorrhizal association with respect to the number of plant species that form them • Found in species in all divisions of terrestrial plants – widely distributed in annuals, perennials, temperate and tropical trees, crop and wild plants • Estimated to occur on 300,000 plant spp.
  • 25. Arbuscular mycorrhizal fungi • All are in the Zygomycota in the Glomales – or newly proposed phylum Glomeromycota • Include ca 130 species in 6 genera • All are obligate biotrophs • Form large spores that superficially resemble zygospores, but not formed from fusion of gametangia – azygospores or chlamydospores • Spore diameters range from 50 to 400 μm
  • 27. Arbuscular mycorrhizae Fossils of spores found that are as old as first land plants – 460 mya
  • 28. Specificity • Few species of fungi and many species of plants – very low specificity • One fungal species may form association with many different plant species • Much different than biotrophic parasites that have a limited host range
  • 29. Morphology • Root morphology is not modified • To detect, must clear and stain root to observe fungal structures • Fungi form both intercellular and intracellular hyphae • Intracellular hyphae analogous to haustoria – called arbuscules – tree like branching pattern • Thought to be site of nutrient exchange between fungus and plant
  • 30. Arbuscules • Surrounded by plant cell membrane • Typically disintegrate after ca 2 weeks in plant cell and release nutrients • Thought to be site of nutrient exchange
  • 31. Vesicles • Intercellular hyphae may also form large swellings – vesicles – at ends of hyphae or intercalary • Typically rich in lipids & thought to be involved in storage
  • 32. AM
  • 33. Arbuscular mycorrhizae • Not as well characterized as ectomycorrhizae • Root is not altered in morphology – difficult to determine when roots are infected – must clear and stain followed by microscopic examination • Fungi are obligate biotrophs – cannot be grown in axenic culture – so difficult to conduct experiments
  • 34. Interaction • Fungus receives organic nutrition from plant – since they are biotrophs, don’t know what their requirements are • Fungus produces extramatrical hyphae that take up inorganic nutrients from soil – particularly P, may also supply N as they may produce proteinases • Increase drought tolerance – many common desert plants are heavily mycorrhizal • May also increase resistance to root pathogens
  • 35. Effect of AM • Growth of plants that are infected better – particularly if soil is poor in nutrients
  • 36. Other types of mycorrhizae • Orchids – orchid seeds are very small and do not contain enough organic reserves to allow development of the plant • Must be infected soon after germination – fungus provides seedling with carbohydrates • Basidiomycetes involved in this mycorrhiza are litter decomposing species of Rhizoctonia, Armillaria that produce cellulases
  • 37. Orchid mycorrhizae • Fungi are widely distributed outside the symbiosis – some are plant pathogens, others are saprotrophs • Appears to be a delicate balance between plant and fungus • Orchid keeps fungus in check by digesting intracellular hyphal coils, production of antifungal substances so fungus doesn’t kill the orchid
  • 38. Orchid mycorrhizae • Not clear about benefits to fungus – may obtain amino acids and vitamins from orchid
  • 39. Ericoid mycorrhizae • Plants are Ericaceae – Erica, Vaccinium - heathland plants • Fungi are Ascomycota and Deuteromycota • Form loose network on surface & hyphal coils inside epidermal cells of hair roots where nutrient exchange is thought to take place • Shown to supply N to plant – fungi secrete proteinases
  • 40. Arbutoid mycorrhizae • Plant are also Ericaceae – Arbutus, Arctostaphylose, Pyrola • Fungi are basidiomycetes that also form ectomycorrhizae • Fungi form sheath and Hartig net, hyphae also penetrate outer coritcal cells
  • 41. Monotropoid mycorrhizae • Plants are nonchlorophyllous – Monotropa • Fungi are basidiomycetes – boletes that form ectomycorrhizae with other plants (conifers) • Plant depends on its mycorrhizal fungus - for its organic nutrients as well as inorganic nutrients
  • 42. Mycorrhizae • Key components of ecosystems • Link plants within a habitat • Labelled CO2 fed to tree can be found in seedlings growing nearby • Retain and conserve mineral nutrients