GLOMEROMYCOTA

1. Symbiosis
2. Roots vs mycorrhizas
3. Arbuscular mycorrhizas

The structures grow in the the cell

BACKGROUND TO SYMBIOSIS:
de Bary: working on lichens and figured out the life cycle of cereal rust: figured out
what causes blight in potatoes; he came with the notion of symbiosis = the living
together of unlike organisms; a continuum from parasitism to mutualism; sometimes
restricted to a persistent mutualism; nowadays, people often refer to the narrow
definition
Pasteur: Microbes produced from preexisting microbes
Petri: Artificial media for axenic culture of microbes
Koch he figured out that anthrax was responsible for tuberculosis; he came of with
the postulates which help us to recognise a microbe:
Presence in hosts
Axenic isolation: isolate and grow them outside the cell
Re-inoculation: find the symptoms again
Re-isolation: be able to isolate/separate them again
the last 3 could not always be true because some organisms cannot de grown
independently

DEFINITIONS:
host: larger and/or evolutionarily older than symbiont; in some cases the size does not
help
parasite: grows at expense of host; negative for the host; sometimes it is very small,
not noticeable, but it still is there
pathogen: interferes with essential host functions, causing significant problems
biotroph: grows and reproduce n or on a living host; often obligate and specific; they
keep their host alive;
necrotroph: thrive on dead host; destructive interaction; you can be a biotroph and kill
your host and then turn into a necrotroph
saprotroph: free-living recycler
synergy: doing something together than neither can do alone; function and/or
symbiosome; pretty common in lichens;

MYCORRHIZAS:
Comparing the ability of plant and fungi to interact with the soil;
Root hairs: small cells that dont branch, non-fusing, unicellular; there are pores,
cracks in the soil where the biological activity takes place; when there is water in
them, there is biological activity; water availability changes over time, so plants/roots
have to cope with that;
Fungi filaments are 10 times thinner than root hairs; they can grow much longer;
branching, fusing and multicellular; the fungi win in a drying soil because they can get
to resources in smaller places; so, fungi will win in soil
Its conveniant to combine plants with fungi
Plants have photosynthesis so they can make carbon to grow in the soil;
An ancient plant-fungal symbiosis; typically mutualistic and obligate; based on
exchange of photosynthesis for soil minerals; involving the majority(90%) of plants
and a minority of fungi

ARBUSCULAR MYCORRHIZAS
In living plant cell, these fungi will make these tree-like
structures
Considered an ancestral condition in land plants (plants
rely on this)
These fungi are unable to grow without the association
Huge spores
Might be the most abundant fungi
Relatively small number of species (cca. 200 species); this
might have something to do with the fact that they are
asexuals
Facilitated plant terrestrialization
In the Rhynie chert fossils, there are structures that look
like this (part of the evidence for their ancestry)

ARBUSCULAE AND COILS:

Very intimate, intracellular interactions
2 types of structures: coil or branch, tree like
the plant cell is still alive: the fungi broke its cell membrane, but did not get into the
cytoplasm, damaging it (the cell cytoplasm will follow the form of the fungi)
increased exchange surface area
the surface area is lined with transporters: trans membrane proteins for P and C
transport
100% of plant P via fungi
100% of fungal C from plant
the fungal structure holds the nucleus; the vacuole will support the tree structure; the
fungi structures go in and out of the cell

MYCORRHIZAL NETWORKS
Fungi are growing n and out of the roots of a plant and growing out in the soil, being
effective in mining the soil for P
They might cross another plant and grow in it, too
So, they connect plants ; they transport resources between plants of different species

MICROSPORIDIA
(ca 1,300 species described so far)
studies by Pasteur
true fungi, possibly derived from the
Chytridiomycota
massive evolutionary reduction: lost almost all
the traits associated with the fungi; they
acquired a set of unique traits found only in this
group
one of the smallest genome known (>2Mb);
no mitochondria or peroxisomes
they are intracellular obligate parasites;
cause chronic debilitation of host; very common
on fish and insects; animals and even humans
they are associated with decline in fungi
the most unique trait is the way in which they deliver themselves in the cell: the most
sophisticated method: inject themselves into the host
enter the gut cell to form xenoma (symbiosome)
both sexual and asexual;
they have horizontal and vertical transmission
intracytoplasmic

outside: 2 cell walls so they are well defended from the nvironment; they can wait
years before being ingested
they can create a lot of pressure within the cell
insisde there is a coil that loops around the cell (inside) = 100 times longer than the
cell;
the vacuole generates pressure, increasing in volume, pushing the cell contesnts; the
tube is pushed out until the cytoplasm of the cell is pushed out through the tube and
it will eventually get into a cell
this is the process of high sped injection
BASIDIOMYCOTA
1. Characteristics
2. Development
3. Biomass recycling (particularly wood)
4. Rusts

Some of the most diverse in shape and colours, comparable with the diversity in
flowers
Second most diverse fungal phylum; over 30Kspeciesdescribed so far
Mainly sapotrophic (degrade something in the environment for their benefit) or even
pathogenic
Spend most of their life as dikaryotic (one nucleus is maternal and one is paternal)
septate filaments
Meiotic basidiospores formed outside a hypha called basidium
There are 4 spores (suggestion of meiotic process)

In between2 adjacent cells: more protected than in ascomycetes; cells are more
distinct, separated than the ascomycetes; there is a cup like structure through which
the cells are connected =dolipore septum; when one cell is damaged, the structure is
occluded with proteins

DEVELOPMENT OF BASIDIA WITH BASIDIOSPORES (MEIOTIC SPORES)

The organism has enough resources so it wants to make some spores
After a karyogamy results a diploid cell (n)
The cell is now going through 2 meiosis, getting a 4 haploids nuclei structure =
sterigma = horns at the top of the filament; the horns are going to develop as
basidiospores; inside of the mother cell a vacuole will start growing, pushing all the
nuclei in the basidiospores which are ready for release;

DIKARYOTIC HYPHAE & CLAMP CONNECTIONS:

Between any 2 consecutive cells, there is a hook structure- = clamp connection
Ensures that in every cell there is a mother nuclei and a father nuclei
As the clamp start to grow backwards, the cell goes through a mitosis: 2 nuclei will
migrate in the top cell (the one that will develop the basidiospores; 1 will migrate
backwards and 1 will get trapped in the clamp connection;
Asexual fussion of 2 filaments:1 new apical cell with 2 nuclei and 1 basic old cell with
2 nuclei;

How to release the spores into the environment:

Sterile supportive tissues and gills that have spores. There are structures that look lie
gills or teeth, pores or some sticky strucutres that allow insects to go in and take
spores

Spores are released in a huge number most prolific organisms; if the cite isremoved
and put it on paper and cover with glass (to be moisturised), the spores will be falling
down;
1 side of one gill and 1 side of other gill; the basidia will mature in the space in
between; it is probably that the spores will get stucked on the basidia so, a
mechanism to push them is needed described by Buller: the spores will go out less
than 1mm and will fall down on the ground ( a lot of moisture in the air); in one of the
spore: there is a hoen structure at one end; a sugar (compound that is hygroscopic) is
secresed and will attract water from the atmosphere; when the tension is too much,
the water will break and will give the spore a push = bullet mechanism; considered
the fastest flights in nature: at the beginning there is 10,000 G, but it slows down
quickly;

LIFE CYCLE:
Sexual life cycle; there are no asexual spores produced;
In the basidium are made basidiospores and Buller force to propel the spore out;
haploid basidiospores are in the environment; if they are lucky enough, they are being
to be able to germinate and will go plasmogamy (will not grow as haploids too long);
n+n state is favoured in the life cycle; there are 2 mating loci that cannot share alleles
they will control the life cycle heterodimeric.
Each cell will have 2 nuclei and will form a mycelium where there are
interconnections; if the asexual fussion is tobe successful, the alleles must be shared;
The fungi will start to make a reproductive structure (agaric) (very variable, at least as
variable as flowers); the terminal cells will start to become basidia and will be, for a
short time diploid (2n) and then will go through meiosis

BIOMASS RECYCLING:
95% of terrestrial biomass is wood
main componnets of wood: cellulose (glucose polymer) cellulases
lignin: complex heteropolymer irregular cross-linking; aromatic
ring backbone; masivelly C-rich and N-poor; insoluble
hemicellulose hemicellulases
pectin (galacturonic acid polymer) pectinases
there is no enzyme that breaks down lignin that traps the cellulose that easily
degradable

white rot fungi break down lignin, revealing cellulose, hemicellulose and pectin;
(saprotrophs and necrotrophs): only some basidiomycetes and a few ascomycetes
but they are not going to be able to use it, but get access to the easy sugars
 they evolved a new different set of enzymes to deal with lignin: not hydrolytic, but
OXIDATIVE EXONZYMES: peroxidases and laccases they carry our Fenton;s reaction
(the most potent oxidising reaction): H2)2 + Fe2+ Fe 3+ + HO- + HO

they must evolve a variety of ways in which they are not attacked by the hydroxyl free
radical themselves compartmentalisation and translocation(for which they evolved
vasculature: dead cells that can transport solutes)

this process leaves something behind; its never degrading everything that finds
behind; without this process, plants will just block in all the C in the atmosphere

RUSTS:
Group of basidiomycetes; ca 9,000 spp of obligate biotrophic parasites described so
far

Puccinia graminis stem rust (black rust; black stem rust)

- Main disease of wheat (Triticum, a monocot): damaged stems that break down
the plant, releasing spores
- A disease of barberry (Berberis, a dicot): on one side of the leaf, it makes 1 kind
of spores and 1 kind on the other side of the leaf;
DE BARRIS: the spores put on healthy wheat plants will do nothing; this is because
they needed an intermediate host;
The basidiospores are not able to infect wheat, but they infect barberry; uredospores
can infect wheat;
The telospores can infect barberry

Cause damage to the host; they are not able to grow without the host;
Weve learnt a lot about plant immunity
from rusts

They are not just typically biotrophic

they depend on 2 hosts that are distantly
related (a monocot and a dicot; a conifer
or a dicot); in order to cpmplete the
lifecycle, they have to alternate between
the 2 hosts
They have to take full advantage of their
life cycle --. 5 different spores types in 1
year
Sometimes they can use insects to move
spores to another plant; sometimes they
sterilise a plant and make pseudoflowers
in order to manipulate insects;
Scent, colour and sugars for insect
dispersal

Repetitive ballistospory and basidiospores

If a spores lands on a plant that is not the right host; the spore, however, can make
another spore and shoot it further to find the right spore;
Sometimes it can use a plant for protection reservoir; the intermediate hosts are not
vectors;
Some rusts have lost it and they have only 1 host

OOMYCETES AND OTHER PSEUDOFUNGI

(early branching EK, which have, more or less, converged on a fungal form of
life)

AMOEBOZOA (more or less terrestrial) might be more closely related to Fungi

and Animals than thought
- Myxomycota
- Plamodiophoromycota
- Acrasiomycota
STRAMENOPILES (more or less aquatic; they need free water in their life cycle)
- Oomycota
- Labyrinthulomycota
- Hyphochytridioycota
MYXOMYCOTA = slime moulds
(cca 900 spp)

probably derived from fungi-animals lineage

they are filamentous
live mostly as free-living haploid amoebe, some with flagella
phatotrophic predators of protists, bacteria and yeasts: they engulf an unicellular
organism in a vacuole
this goes from a unicellular to a multicellular stage
social cooperation, kin recognition and altruism (they give up their own reproduction
to allow others)

DICTYOSTELIUM =model for EK cytoskeleton; unicellular; they are in the environment

and might sense that the resources are scarce so they start to converge, to
aggregate and fuse wit each other. They will move together toward a concentration
gradient;
They fuse only if they have matching alleles; they can only move together now
(plasmodium); they can be in slime mode; once they get resources they form a stock
and a round structure at the top; the cells at the top are going to contribute to
reproduction; the ones at the bottom are going to sacrifice themselves (forming stalk)
apoptosis (so they can be more rigid;
Asexual fusion; they must be closely related, but there can be some differences
Some lineages of cells have undergone selection to place themselves on the top
unresolved problems with multicelullarity cheating cells

OOMYCOTA: filamentous, very fungal looking;

Every organism has a disease caused by one of these
Most grow as aseptate diplodid hyphae (multicellular)
Cellulose cell walls: this normally happens in plants
Flagellated spores
Aquatic(free living and typically selfers - inbreed) or terrestrial(much finer filaments,
typically biotrophic, outcrossing) change in lifestyle
Saprotrophs, facultative necrotrophs or obligate biotrophi haustorial parasites (put a
filament in the host, but will keep it alive) (aggressive)
Main pathogens of potatoes, tomatoes and grapes (losses >2.5 billion/yr)
Great Famine decimated the potato crop across Northern Europe
Sudden Tree Death

LIFE CYCLE (Phytophthora infestans) Variety of ways of infecting hosts

A filament(aseptate) with diploid nuclei (2n) arrived from American on a ship in 1845
with extreme rate of spread;
It grows into the shoots of the plants; they come out of the stomata and produce
reproductive structures lemon shape kind of structures = sporangia (2n) the
drops will form around the sporanfiospores and they will get trapped in a dew drop
which will catapult the sporangia out;
Now that they are out, they have some options; if its warm and dry, they will form
filaments that will grow into stomata (come out of the stomata and go back into it) ->
they form an apersorium that will help them to push through; this is an asexual cycles

If its cooler and there is free water; instead of germinating directly, they can
differentiate into zoospores (heterokonts, 2 kinds of flagella: cilli- tinsel, rigid and a
whiplash which will allow movement); it will swim around and when finds a leaf, it will
produce a cell wall and become sticky, forming a cyst; now they will germinate and
generate and a filament that enters stomata (still asexual)

1981 (mat A1): is another mating type and now they are going to go through a sexual
cycle; they need to have the same mating types in the same plant; they will release
pheromones so that they can find each other;
they will go through a fusion: one filament is going to grow (male) through the other
one(female); this is the only point of the life cycle when a septae is going to be made;
the terminal cells will go through meiosis (resulting 4 diploid nuclei) and then 3 are
going to be degraded), ending up with 1 haploid nucleus in each cell; the male
nucleus is going to migrate in the female and will end up with 2 haploid one; a new
cell is going to be formed in the female cell = oospore; it will be a dikaryon during
winter;
the diploid is the favoured state; a oospore that has germinated has a tube and has a
diploid structure that can be released in the environment; they can either generate a
micellium with filaments or, if conditions are wet and cool, the flagella will allow them
to move around and find plants they need;

even when it goes through asexual reproduction, they still can increase genetic
diversity a part of the genome is full of transposable elements (involved in infecting
the hosts)