Fungi are organisms with a common lifestyle

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Storage compounds – retaining nutrients

Littlefield and Heath 1979

Ultrastructure of Rust Fungi lipid glycogen

Nutrition of biotrophs

• Components are extracted through haustoria

• Nutrients are soluble and organic

• Extracellular degradation for cell penetration

• Extracellular factors establish/maintain a compatible infection

• Suppress senescence

Suppressing senescence www.mpiz-koeln.mpg.de/schlef/PSL_webpage.html

Livning substrates exploited by fungi

What is the nutrient flow direction?

http://www.ucmp.berkeley.edu/fungi/rhyniefungus.jpg

Arbuscular and ectomycorrhizal fungi

Amino acid biosynthesis

Secondary metabolites

• Glucose-derived – polysaccharides, peptidopolysaccharides, and sugar alcohols.

• Condensation products of acetate – derived from the acetate-malonate pathway of fatty acid synthesis, e.g. polyketides and phenolics.

• Condensation products of acetate derived from the mevalonic acid pathway, e.g. terpenes.

• Phenolics derived from the shikimic acid pathway of aromatic amino acid synthesis.

• Derivatives of other amino acid syntheses.

Secondary metabolites

Pigments

Hormones

Toxins

Co-regulated with sporulation

Secondary metabolites of Saccharomyces www.crc.dk/flab/ newpage13.htm

Genetics – study of heredity

• Transmission - the passage of traits from one generation to the next

Genetics – study of heredity

• Population - genetic diversity and change within natural populations

Genetics – study of heredity

• Molecular - details of gene structure and function

Our focus for genetics

• transmission and molecular genetics in experimental systems

• defining a population

– organisms in culture

– humungous fungus

– vegetative incompatibility

Transmission genetics

• Typical characteristics of fungal genomes

Small

– S. cerevisiae 6 MB – 6000 genes

– A. nidulans 13 MB – 12000 genes

– H. sapiens 1300 MB – 30000 genes

Typical characteristics of fungal genomes

• Little repetitive DNA – single copy genes

– 50-60% of nuclear genome is transcribed into mRNA in

S cerevisiae

– 33% in S. commune (basidiomycete)

– 1% in humans

• Introns

– few, often none

– small – 50-200bp vs ≥10 kb in mammals

Most higher fungi are vegetative haploids

• One genome copy per nucleus

• Alternatives?

– Plants?

– Algae?

– Animals?

Risks of haploidy

• No backup copy in case of genetic damage from UV or chemical mutagens

• Yeasts tend to be diploid ( S. cerevisiae except for lab strains) or have short G1 (S. pombe )

Chant and Pringle JCB 129:751

Advantages of haploidy

• A multinucleate cell can expose genome to mutagens

– most mutations are deleterious

– select for advantageous mutations in a heterokaryotic system

• Phenotypes of recessive mutations are obvious in the vegetative state, without generating homozygous recessives

• Lab strains of S. cerevisiae now generally include a mutation which stabilizes the haploid state

Transmission genetics – passage of inheritance

• Similar to more familiar mammalian systems, with bulk of life cycle haploid

• ‘Genders' are ‘mating types’

– cells are biochemically distinct but morphologically identical

Fungal mating systems

No mating factors  A. nidulans

• Inbreeding possible

– disadvantage – sex does not necessarily increase genetic diversity

– advantage – can form resistant spores even if no mating partner is available

– A. nidulans ascospores from 1995 still viable after 4°C storage, whereas conidia viability is severely reduced after several months at 4°C

One factor (zygo, asco, some basids)

• Bipolar mating system

• meiosis will give two types of segregants

– N. crassa a and 

Rhizopus + and

One factor (zygo, asco, some basids)

Advantage

– outbreeding

Disadvantage

– cannot produce resistant sexual spores unless a partner is available

• ‘Coping’ with one-factor mating systems

Some fungi have multiple alleles at the mating locus

Mating type switching in Saccharomyces

One factor (zygo, asco, some basids)

• In

S. cerevisiae "a" cells produce a-factor, a peptide sexual hormone, and

-receptor; converse for

 cells

• hormones/receptors interaction promotes schmooing, wall changes promote adhesion

Two factors, A/B (often in basids)

• Tetrapolar mating system  meiosis give four types of segregants

• A

1

B

1

:: A

2

B

2

A

1

B

1

, A

1

B

2

, A

2

B

1

, A

2

B

2

A and B functions are distinct

• in homobasids (.....?)

– A controls pairing and synchronous division of nuclei, hook cell formation;

– B controls septal dissolution and hook cell fusion ( b -glucanase activity) and nuclear migration

A and B functions are distinct

• in heterobasids (....?)

– A controls pathogenicity;

– B controls filamentous growth

Systems restricting outcrossing in one-factor mating type systems

• self-fertility  S. cerevisiae has "mating type switching"

• molecular basis  both mating genes have a storage site and an expression site.

• if the appropriate partner cell is not available when mating conditions are presented (how would this be detected?)will induce swi expression

Systems restricting outcrossing in one-factor mating type systems

• vegetative (somatic) incompatibility

• het genes are important for mating, but prevent vegetative fusion

Systems restricting outcrossing in one-factor mating type systems

• vegetative (somatic) incompatibility

• in Fusarium – vegetative incompatibility is important for maintaining distinct populations with different host specificities

• Fusarium oxysporum f. sp. groups

Mutants in experimental fungal systems

• spontaneous mutations or mutagenesis (uv, chemicals)

• each gene is named for 1st described mutation

• Example: gene for pigmentation is called

“white” because the mutant lacked colouration

Different species, different naming system

• Saccharomyces cerevisiae

• Schizosaccharomyces pombe

• Aspergillus nidulans

• Neurospora crassa

• Generally, three-letters plus a letter or number – hypA, CDC2, cdc28

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