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Mycorrhizas

Fungal Symbiosis

Literally means fungus root

Fungi form many types of symbio8c
associa8ons
Roots
Underground organs of plants

Mutualis8c associa8ons
Cassie Doino, Andreal Reed, Ka8e
Allen, and Adie Kurtanich

Types of Mycorrhiza

Arbuscular mycorrhizas
Ectomycorrhizas
Ericoid mycorrhizas
Orchid mycorrhizas
Monotropoid mycorrhizas

Fungus sends nutrients to plant

Plant provides fungus with sugar

Arbuscular Mycorrhiza (AM)

Most common type of mycorrhiza
Found worldwide
Crop plants
Trees
Herbaceous plants

Originally classied as Zygomycota

Now classied in new monophyle8c group
Glomeromycota

Arbuscular Mycorrhiza (AM)

Roots show no outward signs of infec8on
Hyphae can only be seen by special
techniques
Dieren8al interference contrast microscopy
Staining

Arbuscular Mycorrhiza (AM)

Fungal hyphae extensively colonize roots
1. Grow between root cor8cal cells
Produce large swollen vesicles
Storage func8on

Treat root with strong alkali

Stain with fungal dyes such as trypan blue

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Arbuscular Mycorrhiza (AM)

Fig 13.1

Fungal hyphae extensively colonize roots

2. Penetrate individual root cor8cal cells
Form tree-like branches known as arbuscules
Main site of nutrient exchange between fungus and cell root
Remain alive for short period of 8me
Degenerated and replaced

Fig. 13.1. The principal features of arbuscular mycorrhizal (AM) fungi, observed by clearing the root 8ssues with strong alkali and then staining
roots with the fungal dye, trypan blue. Top le>: A root heavily colonised by AM fungi, with hyphae that radiate into the soil. Top right: When
observed through the depth of the root cortex, AM fungal hyphae are o^en seen to run parallel to the root axis, growing between the root
cor8cal cells. These hyphae are irregular, with constric8ons and bulges, quite unlike the hyphae of most other fungi. They frequently produce
large, swollen vesicles within the root 8ssues. BoBom le>: some of the external hyphae and hyphal aggregates produce clusters of spores in the
soil. BoBom right: Some of the root cor8cal cells are penetrated by hyphae that branch repeatedly to produce intricately branched arbuscules,
o^en completely lling the root cells. [ Jim Deacon]

Arbuscular Mycorrhiza (AM)

Spores
Large, up to 400 um
Germinate and infect the roots
From an appressorium-like infec8on structure on the
root surface

Arbuscular Mycorrhiza (AM)

Arbuscular Mycorrhiza (AM)

Ecological Signicance of AM
AM fungi provides plants with mineral nutrients
from soil
Especially phosphorous
Produce extensive hyphal networks in soil
Release acid phosphatases
Absorb phosphate in excess and store it in form of
polyphosphates

Fig 13.4

Host ranges and communi8es of AM fungi

AM have a wide range of host
Experimental study by Van den Heijden et al.
Plants in natural communi8es are colonized
preferen8ally by dierent strains of AM fungi
The diversity of AM fungi in a site can inuence the
plant biodiversity in natural ecosytems

Fig. 13.4. An increase in the number of dierent arbuscular mycorrhizal fungi in a soil leads to an increase in plant
produc8vity and plant biodiversity.

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Fig 13.5

Ectomycorrhizas
Found mainly on woody plants
Coniferous and broad-leaved trees outside of
tropics
Such as pine, spruce, oak, beech, and birch

Members of Basidiomycota or Ascomycota

Characteris8c feature
Presence of substan8al sheath of fungal 8ssue
Incases the terminal nutrient absorbing rootlets

Fig. 13.5. Total biomass and the biomass of three representa8ve plant species grown in soil with no mycorrhizal fungus (0) or with
four separate AM fungal species (A,B,C,D) or a combina8on of all four AM species. Note that the ver8cal scale of each histogram is
dierent but the largest biomass is shown in each case.

Ectomycorrhizas
Ecologically adapted to grow as symbionts
Can grow in laboratory on simple media
Have lifle to no ability to degrade cellulose and
lignin

Rarely have high degree of host-specicity

Ectomycorrhizas
Grouped into two types
Mature
Host restricted
Mature trees
Release proteases
Providing ectomycorrhiza plants with nitrogen in the form of
amino acids

Ectomycorrhizas
Grouped into two types
Generalists
Wide host range
Young trees
Such as Laccaria, Hebeloma, Thelephora terrestris

Ectomycorrhizas
Fungal Networking
Extensive network of hyphae and mycelial cords
Link dierent plants within a habitat
Mycelial connec8ons retain mineral nutrients
Withdraws nutrients from degenera8ng mycorrhizas
Transports nutrients to other func8oning mycorrhizas

Fungal network extends beyond root zone

Mycelial cords transport water from deeper in the soil, beyond
the reach of the roots

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Fig 13.7

Fig 13.8

Fig. 13.7. Le>: Scanning electron micrograph of a cross sec8on of part of a mycorrhizal root, showing the fungal sheath that
surrounds the root. Right: Thin sec8on of part of an ectomycorrhizal root. The arrowheads show hyphae invading between the root
cor8cal cells, forming the Har8g net. Nutrient-exchange between the fungus and the root is thought to occur in this interfacial
region.

Figure 13.8 A young larch seedling, about 3 cm high, growing in a peatbased substrate
against a sloping face of an observa8on chamber. Mycoohizas can be seen at the base
of the stem but almost all the visible growth is mycelial cords that explore the soil for
nutrients.

Ericoid Mycorrhizas

Fig 13.10

Typical host families

Ericaceae
Epacridaceae
Both families produce coils of hyphae termed hair
roots

Fungi that produce this mycorrhiza are free-

living saprotrophs in soil
Ascomycota
Fig. 13.10. Sec8on through part of the protocorm (basal stem region) of an orchid, Neo4a, showing coils of hyphae (termed
'peletons') within the orchid cells.The plant cells were alive at the 8me of sec8oning, evidenced by presence of nuclei (the dark
granular structures) in two of the orchid cells. [ Jim Deacon]

Ericoid Mycorrhizas
Grown in culture
Produce septate hyphae with fragmented zigzag
growth

Primary role
To provide host plants with nitrogen

Secrete proteinase
Release amino acids from soil organic mafer

Orchid Mycorrhizas
Parasi8c on fungus
Orchid seeds
Small, consists of embryo and few nutrient
reserves
Triggered to germinate
Produce few root hairs
Must be colonized by a fungus at an early stage or
seedling will die

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Orchid Mycorrhizas
Fungus penetrates orchid embryo and produces
hyphal coils
Peletons
Only last a few days
Degenerate and replaced by further coils in other cells

Repeated process provides main source of

sugar to developing orchid

Monotropoid Mycorrhizas
Plants of family Monotropaceae lack
chlorophyll
Dependent on mycorrhiza fungi for all their
nutrients

Found in shade beneath forest trees

Fungi involved are Basidiomycota

Orchid Mycorrhizas
Provide orchids with sole source of
carbohydrates during early years of life
Most do not emerge above ground or produce
chlorophyll un8l 3-5 years old
As many as 200 species never produce chlorophyll
Remain dependent on mycorrhizal fungus

Monotropoid Mycorrhizas
Three membered symbiosis
Direct nutri8onal connec8on
Tree host
Mycorrhizal fungus
Parasi8c higher plant

Radiate into the soil as hyphal networks or

mycelial cords
Form hyphal sheath around roots

Summary

Lichens

Mycorrhizal type

Typical host plants Fungi involved

Major Signicance

Arbucular
mycorrhizas

Many

Glomeromycota

Phosphorous
uptake from soil

Ectomycorrhizas

Forest trees

Basidiomycota,
Ascomycota

Nitrogen uptake
from soil

Ericoid mycorrhizas Heathland plants

Ascomycota

Nitrogen uptake
from soil

Orchid mycorrhizas Orchids

Basidiomycota

Fungi supply plant

with sugars

Monotropoid
mycorrhizas

Nonphotosynthe8c Basidiomycota
plants

Plants obtain sugars

from
ectomycorrhizas

Lichens symbio8c associa8ons of fungus &

photosynthe8c partner (green alga or
cyanobacterium)
Involve 2 or more partners
Come together to form dis8nct new organism
Form long-term symbio8c rela8onships

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Lichen Partners
13,500 to 17,000 species of lichens
Dicult to name & classify since composed of
2 or more organisms
Mycobiont fungal partner
Usually type of Ascomycota or rarely
Basidiomycota

Photobiont photosynthe8c partner

Green alga or cyanobacterium
Examples of Lichens

Fungal Lichen Partners

Lichen fungi are very specialized & mainly
found only in lichen partnerships
Few species are found in non-symbio8c states
Some species have been grown in lab
Slow growth
Lack enzymes needed to live freely

Algal Lichen Partners

About 100 types of green algae are lichen
partners
Trebouxia
Most common genus associated with lichen
partnerships
Single-celled rarely found as free-living
Found in temperate regions

Trentepohlia
Found in tropical regions

Cyanobacteria Lichen Partners

Found in about 10% of lichens
Pel9gera
Most common genus
Found in dog lichen

Some lichens have green algae & cyanobacteria

Cephalodia wart-like structures on lichen
surface
Thought to use nitrogen xing abili8es of
cyanobacteria

Forms of Lichens
Lichen thallus body of lichen
Categories
Foliose
Flat & leaf-like

Fru8cose
Erect & branch-like

Squamulose
Small & scale-like

Crustose
Flat & crust-like

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Structural OrganizaUon
Many have well-dened structures
Typical structure
Upper cortex
Medulla
Lower cortex
Rhizinae

Algal associa8ons also require gas exchange

Physiology: Water Requirements

Lack of water
Lichens containing green algae absorb water from
humid air & resume photosynthesis
Lichens containing cyanobacteria must absorb
liquid water to resume photosynthesis
Drought tolerance is likely conferred by
hydrophobins produced by lichen fungi

Hydrophobin coated fungal hyphae create spaces

for gas exchange

Physiology: Nutrient Exchange

Physiology: Nutrient Exchange

Lichen fungi protect photosynthe8c partner &

absorb nutrients

Photosynthe8c partners provide sugars for the

growth of lichen

Very ecient at absorbing nutrients from limited

environmental resources
Unfortunately for lichens, also absorb toxins &
pollutants
Thus, lichens are rarely found in polluted environments

ReproducUon & Dispersal

Propaga8on
In some lichens, fragments break o & are
transported by wind or animals
Isidia stalk-like structures produced by some
lichens that break o
Soredia powdery propagules
Dispersal of fungal ascospores & reassembly with
photosynthe8c partner

Give up to 90% of photosynthate to fungal partner

Carbohydrates found in lichens

Mannitol
Arabitol
Ribitol
Glucose

Ecology & Signicance

Lichens can live in a variety of environments
From tundra to cooled lava ows to tropical
forests

Most signicant role of lichens

Contribu8on to soil forma8on

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Summary
Symbio8c associa8ons between fungal,
cyanobacterial &/or algal partners
Can survive in many environments
Have dened structures
Have several ways to propagate
Signicant contribu8on to soil forma8on

Geosiphon pysiform

Geosiphon pysiform
Unique rela8onship
between mycorrhizal
and cyanobacteria
Duel organism

Discovered in Germany
1996

Belongs to the
arbuscular mycorrhizal
(AM) and related fungi

Bladders of Geosiphon pyriforme growing

on the surface of soil.
Courtesy of A. Schuessler

Geosiphon pysiform

Endocytose cyanobacteria
Leads to a membrane bound symbiosome containing
cyanobacteria

Provides fungus with source of sugar

Symbio8c

Cyanobacteria depends on phosphate from

fungus
Mutualis8c
Image courtesy of A. Scheussler & M. Kluge, 2001

Fungus/Insect Mutualism
Several insects have mutalis8c associa8ons
with fungi
Provide the insect with a food source
Insect ensures mutualism

Carrying and transmiqng the fungus

Mycangia

Inoculate suitable substrate with spores on the

fungus
Tending the substrate to promote growth of
fungus
Food source

Sirex Wood Wasp

Female injects toxic mucus and Amylostereum
areolatum
Laying eggs in bark of damaged trees
Mucus kills tree cells
Fungus feeds on the killed wood
Insect larva feed on fungus

A^er pupa8on, the female adults collect

the spores from the roqng wood
Store in mycangia to repeat the cycle

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Leaf-Cu^ng Ants
Aqne Ants

Produce large nests

Carries fungus, Leucoagricus, in pouch of mouth
Deposits fungus on suitable plant
Eggs laid as fungus grows
Worker ants bring leaf pieces back to nest, inoculate
with fungus
Fungus produces hyphae

The wood wasp, Sirex noc9lio, boring a hole in a weakened tree to deposit eggs
and fungal spores.
Courtesy of M. P. CouDs, J. E. Dolezal and the University of Tasmania

Gardening Termites
Do not digest wood
Evolved symbiosis with the Termitomyces
Found in fungus gardens

Termites weed out any contaminant fungi

Heavy rainfall
Fungus produces large, mushroom fruitbodies

Gongylidia or broma8a
Fed to larvae
Provide enzymes needed for degrading plants

Wood Boring Beetle

Produce tunnels in damaged trees
Deposit eggs
Smear with ambrosia fungi

Spores stored in special sacs of beetle

Mycangia

Developing beetle larvae

Create galleries beneath bark
Feed on fungus

Fungus/Insect Mutualism
About 40-60 million years ago, these three
types independently evolved the ability to
culture fungi as a source of food.
Now all of these fungus-cul8va8ng insects and
their fungal partners are mutually dependent
on another.
There is no evidence that they have ever
reverted to an independent existence.

QuesUons
1. Which fungal phylum is associated with
arbuscular mycorrhizas?
a. Zygomycota
b. Ascomycota
c. Basidiomycota
d. Glomeromycota

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QuesUons
2. T/F Ectomycorrhizal fungi are classied as
either mature or immature.
3. T/F The primary role of the ericoid
mycorrhizas are to provide the host plants
with sugar.

QuesUons
5. Lichens that produce at crusts on rock, soil,
or tree surfaces
a. Squamulose
b. Fru8cose
c. Foliose
d. Crustose

QuesUons
4. Stalk-like structures that aid in propaga8on
a. Soredia
b. Cephalodia
c. Isidia
d. Rhizinae

Sources
Deacon, J. Fungal Biology. Cambridge, MA.
Blackwell. 2006.
hfp://www.perspec8ve.com/nature/fungi/
lichens.html
www-biol.paisley.ac.uk/biorefChlorophyta
www3.isrl.uiuc.edu/~openkeyAlgae/image/
Papilla-pore?N=D
hfp://www.lichen.com/vocabulary.html

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