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Growth, Differentiation and Sexuality

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Mating Types in Euascomycetes 295<br />

Fig. 15.1. Life cycle of Euascomycetes. Outer circle: self-compatible life cycle. Inner circle: self-incompatible life cycle<br />

(adapted from a drawing by S.-W. Lu)<br />

III. Description<br />

of Mating-Type Structure<br />

A. General Organization of Mating Types<br />

In all self-incompatible filamentous Ascomycetes<br />

examined to date, the mating-type locus contains<br />

one of two different sequences occupying the same<br />

chromosomal locus in their genome. The extent of<br />

dissimilarity ranges from 1 to almost 6 kb. Metzenberg<br />

<strong>and</strong> Glass used the word “idiomorph” to<br />

denote these large sequences not obviously related<br />

by structure or common descent (Metzenberg <strong>and</strong><br />

Glass 1990). The analysis of self-compatible species<br />

indicates that, in most cases, both mating types are<br />

contained in the haploid genome, usually tightly<br />

linked, but sometimes not, <strong>and</strong> thus these loci cannot<br />

be called idiomorphs, since there is no second<br />

strain of opposite mating type. We retain the use<br />

of the term “mating type”, instead of “idiomorph”,<br />

for self-compatible species. A st<strong>and</strong>ardized nomenclature<br />

for idiomorphs <strong>and</strong> mating-type genes has<br />

been proposed, <strong>and</strong>, in most cases adopted, to<br />

cope with the rapidly growing number of described<br />

mating-type loci from various species (Turgeon<br />

<strong>and</strong> Yoder 2000). This st<strong>and</strong>ardized nomenclature<br />

is based on the constant presence in one idiomorph<br />

of an ORF encoding a protein with a motif called<br />

the α1 domain, found initially in the MATα1 transcription<br />

factor of Saccharomyces cerevisiae. This<br />

idiomorph is called MAT1-1, <strong>and</strong>thecorresponding<br />

gene MAT1-1-1. The other idiomorph is characterized<br />

by the constant presence of an ORF encoding<br />

a protein with a DNA-binding domain of<br />

the high-mobility group (HMG) type <strong>and</strong> is called<br />

MAT1-2. The corresponding gene is called MAT1-<br />

2-1. The number assigned to any particular gene<br />

within an idiomorph corresponds to that of its homolog<br />

in other fungi that have already characterized<br />

MAT genes. If there is no apparent homolog,<br />

the next sequential number is assigned. If a second<br />

mating-type locus is discovered in the same fungus,<br />

it would be MAT2.Thissystemisadheredtobythe<br />

fungal community, with the exception of members<br />

of the Neurospora crassa <strong>and</strong> P. anserina communities,<br />

since designation of genes from these genera<br />

was established before st<strong>and</strong>ard terminology<br />

was proposed. To date, for self-incompatible Ascomycetes,<br />

including the models Cochliobolus heterostrophus,<br />

P. anserina,<strong>and</strong>N. crassa,onlyasingle

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