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
In ectomycorrhizal (ECM)
roots, the fungus forms
extraradical hyphae and a
fungal sheath outside the
root (A) and the Hartig
net surrounding root cells
(B) hiding epidermal cells
and cortical (cc) cells (C).
(A) In fungal cells, the uptake of Pi occurs mostly
through Pht1 phosphate transporters.

To date, only HcPT1.1, HcPT2 (Tatry et al., 2009), and

BePT (Wang et al., 2014) genes have been characterized
by heterologous expression in yeasts.

Genomics and transcriptomic data suggest that other

transporters may play a role in phosphate uptake (e.g.,
HcPT1.2, LbPTs, AmPTs, TvPTs, TmPT3 (Pht2; Casieri
et al., 2013).
(B) In the Hartig net, fungal and plant cells have a common
apoplastic space with no direct symplastic communication.

It is hypothesized that the hydrolysis (a) of polyphosphate

(PolyP) increases Pi concentration in the cytosol of the fungus.

Up to now, the molecular mechanisms sustaining P efflux from

the fungus (b) to the apoplast and P influx (c) from the apoplast
to the plant cell have not been identified.

It is also hypothesized that fungal P transporters may not be

functioning (d)
(C) In plant cells, phosphate ions enter through plant P
transporters. Little is known about plant transporters
responsible for Pi acquisition in ECM roots. Only phosphate
transporters from Populus trichocarpa (PtPTs; Loth-Pereda et
al., 2011) and Eucalyptus marginata (EmPhts; Kariman et al.,
2014) have so far been identified.

Transcriptomic data for Pinus pinaster (Canales et al., 2013)

showed putative encoding sequences for phosphate
transporters (PpPTs).
Full lines indicate transport systems whose
capability in phosphate transport has been
verified by heterologous expression in
yeast.

Dotted lines indicate transport systems

whose involvement in phosphate
transport during mycorrhizal symbioses is
suggested by genomic or transcriptomic
data.
Ed: endodermal cells. Hc: Hebeloma
cylindrosporum, Be: Boletus edulis, Tm:
Tuber melanosporum, Am: Amanita
muscaria, Lb: Laccaria bicolor, Tv:
Tricholoma vaccinum, Pt: Populus
trichocarpa, Em: Eucalyptus marginata,
Pp: Pinus pinaster.
 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.
Restriction Fragment Length Polymorphism (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 14CO2 demonstrated that 14C
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