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)