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Archaeorhizomycetes: a new class for a major clade of ... - IMA Fungus

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RESEARCH News<br />

this involves recognition, directed hyphal<br />

growth leading to conjugation, and then<br />

plasmogamy <strong>of</strong> compatible mating partners.<br />

In order to investigate this complex and<br />

fascinating system, last December Kellner et<br />

A<br />

support values<br />

MrBayes RAxML<br />

< 90 < 50<br />

90-95 50-80<br />

> 95 > 80<br />

0.1<br />

hybrid filament<br />

pheromone response<br />

1 a1 allele<br />

2 a2 allele<br />

3 a3 allele<br />

Ustilaginaceae<br />

host age: 83-89 mya<br />

al. (2011) reported on elegant investigations<br />

designed to illuminate our understanding <strong>of</strong><br />

the evolution <strong>of</strong> the PR system. Ten species<br />

spanning 100 Myr <strong>of</strong> evolution <strong>of</strong> the system<br />

were selected <strong>for</strong> genomic and biological<br />

S. walkeri<br />

S. scitamineum<br />

S. anthracoideisporum<br />

S. mishrae<br />

S. andropogonis<br />

S. reilianum<br />

S. exsertum<br />

S. bursum<br />

U. vetiveriae<br />

U. maydis<br />

U. cynodontis<br />

U. xerochloae<br />

U. hordei<br />

U. strii<strong>for</strong>mis<br />

Me. pennsylvanicum<br />

U. fili<strong>for</strong>mis<br />

U. spermophora<br />

---------------------------------<br />

1 2 3 3 2 3<br />

---------------------------<br />

2 1 1<br />

2 2<br />

2 2<br />

------------------------ 1<br />

1 1<br />

--------------------<br />

1 1 3 3 3 1<br />

----------------------------<br />

3 1<br />

1 3 3<br />

----------------------------<br />

2<br />

1<br />

1 3<br />

study. In addition to detailed comparative<br />

in<strong>for</strong>mation on the alleles, they per<strong>for</strong>med<br />

interspecific sex tests which revealed a high<br />

potential <strong>for</strong> hybridization between species<br />

linked to pheromone signalling. While the<br />

system is optimized <strong>for</strong> within-species sex, it<br />

reveals that there are possibilities <strong>for</strong> hybrid<br />

generation which could lead to smuts with<br />

<strong>new</strong> host specificities. This possibility was<br />

confirmed by the demonstration <strong>of</strong> actual<br />

fusions between not only species <strong>of</strong> the<br />

same genus, but ones in different genera,<br />

as illustrated in the accompanying figure.<br />

The authors comment that the system now<br />

revealed may serve as a valuable model <strong>for</strong><br />

the study <strong>of</strong> the hybrid-based genesis <strong>of</strong><br />

novel genotypes.<br />

Kellner R, Vollmeister E, Feldbrügge M, Begerow D<br />

(2011) Interspecific sex in grass smuts and the<br />

genetic diversity <strong>of</strong> their pheromone-receptor<br />

system. PLoS Genetics 7: e1002436.<br />

S. consanguineum<br />

Ustilaginales<br />

host age: 113-117 mya<br />

pra1<br />

U. maydis<br />

S. reilianum<br />

S. walkeri 80<br />

Me. pennsylvanium 63<br />

U. hordei<br />

* ** *<br />

U. xerochloae<br />

U. cynodontis<br />

Us. gigantosporum<br />

*<br />

B<br />

: bt > 91<br />

0.5<br />

Malassezia pachydermatis<br />

100<br />

100<br />

100<br />

Ma. eriachnes<br />

U. williamsii<br />

Us. gigantosporum<br />

Us. standleyanum<br />

Cintractia limitata<br />

Schizonella melanogramma<br />

Malassezia globosa<br />

U. maydis<br />

S. reilianum<br />

pra2 Ma. eriachnes<br />

61*<br />

U. hordei<br />

82<br />

Us. gigantosporum<br />

71<br />

pra3<br />

S. reilianum<br />

* S. walkeri<br />

U. xerochloae<br />

Us. gigantosporum<br />

Tranzscheliella hypodytes<br />

---------------------------------<br />

1 2 3 3<br />

Melanotaenium euphorbiae<br />

C<br />

Urocystis eranthidis<br />

Figure XXX. Interspecific sex in grass smuts - modified from Kellner et al. 2011 (PLoS Genetics)<br />

(A) Multi-gene phylogeny and interspecific sexual compatibility <strong>of</strong> Ustilaginales. Concatenated Maximum<br />

Likelihood (ML) analysis <strong>of</strong> 2571 bp <strong>of</strong> ssu, ITS, lsu rDNA, ef1-α and rpb1. Circles next to branches indicate<br />

bootstrap support values and a posteriori probabilities <strong>of</strong> Bayesian and ML analyses, respectively. Branch lengths<br />

correspond to substitutions per site and abbreviated branches indicate longer branches. Connected squares<br />

illustrate<br />

It has previously<br />

hybrid filament<br />

been<br />

<strong>for</strong>mation<br />

recognized<br />

(bold lines)<br />

that selfperpetuating<br />

et al., 2005 changes (PNAS). in protein (B) Phylogeny structure <strong>of</strong> mating type-specific <strong>of</strong> the protein pheromone and can receptors. adopt Maximum an amyloid Likelihood<br />

or pheromone<br />

Sup35<br />

response<br />

that<br />

(thin<br />

is not<br />

lines).<br />

essential<br />

Numbers<br />

to<br />

in<br />

the<br />

squares<br />

function<br />

represent<br />

respective a mating types. Coloured boxes depict different phylogenetic <strong>clade</strong>s (see text). Host ages refer to<br />

Prasad<br />

analysis<br />

can be heritable<br />

<strong>of</strong> complete<br />

elements<br />

pheromone<br />

in<br />

receptor-coding<br />

yeasts separate<br />

sequences.<br />

configuration<br />

Numbers and<br />

which<br />

asterisks<br />

self-perpetuates<br />

next to branches<br />

and<br />

indicate<br />

bootstrap (bt) support values and branch lengths correspond to substitutions per site. (C) Interspecific mating <strong>of</strong><br />

from and preceding genetic change (True<br />

& Lindquist 2000); such self-perpetuating<br />

epigenetic structures are termed prions.<br />

Halfmann et al. (2012) investigated the<br />

yeast translation-termination factor prion<br />

haploid sporidia <strong>of</strong> Sporisorium reilianum (Sr) and Sporisorium scitamineum (Ss).<br />

Sr<br />

Ss<br />

1<br />

Interspecific sex in grass smuts (modified from<br />

Kellner et al. 2011). A, Multi-gene phylogeny and<br />

interspecific sexual compatibility <strong>of</strong> Ustilaginales.<br />

Concatenated Maximum Likelihood (ML) analysis<br />

<strong>of</strong> 2571 bp <strong>of</strong> ssu, ITS, lsu rDNA, ef1-a and rpb1.<br />

Circles next to branches indicate bootstrap support<br />

values and a posteriori probabilities <strong>of</strong> Bayesian and<br />

ML analyses, respectively. Branch lengths correspond<br />

to substitutions per site and abbreviated branches<br />

indicate longer branches. Connected squares<br />

illustrate hybrid filament <strong>for</strong>mation (bold lines)<br />

or pheromone response (thin lines). Numbers in<br />

squares represent respective a mating types. Coloured<br />

boxes depict different phylogenetic <strong>clade</strong>s (see text).<br />

Host ages refer to Prasad et al. (Science 310:1177–<br />

1180, 2005. B, Phylogeny <strong>of</strong> mating type-specific<br />

pheromone receptors. Maximum Likelihood analysis<br />

<strong>of</strong> complete pheromone receptor-coding sequences.<br />

Numbers and asterisks next to branches indicate<br />

bootstrap (bt) support values and branch lengths<br />

correspond to substitutions per site. C, Interspecific<br />

mating <strong>of</strong> haploid sporidia <strong>of</strong> Sporisorium reilianum<br />

(Sr) and S. scitamineum (Ss); SEM micrograph.<br />

Prions and phenotypic inheritance in wild yeasts<br />

leads to increased stops in codon readthrough;<br />

that leads to a variety <strong>of</strong> <strong>new</strong> traits.<br />

The prions had been considered an artefact<br />

<strong>of</strong> strains kept in culture, but these authors<br />

examined occurrences and screened <strong>for</strong> <strong>new</strong><br />

prions in around 700 wild Saccharomyces<br />

strains. Prions proved to occur in about<br />

one third <strong>of</strong> the wild strains examined.<br />

Modifications <strong>of</strong> the Sip35 prion were<br />

demonstrated to confer characters likely to<br />

be beneficial to the yeasts under selective<br />

pressures, that is to develop beneficial<br />

phenotypes. Indeed, 40 % <strong>of</strong> the prions in<br />

(30)<br />

<br />

ima fUNGUS

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