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Growth, Differentiation and Sexuality

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Table. 15.2. Structure of the MAT locus in self-compatible Euascomycetes<br />

Mating Types in Euascomycetes 299<br />

Accession number Genes Features<br />

Loculoascomycetes<br />

Cochliobolus homomorphus AF129741 MAT1-2-1::MAT1-1-1 HMG+α domain<br />

Cochliobolus luttrellii AF129740 MAT1-1-1::MAT1-2-1 α domain+HMG<br />

Cochliobolus kusanoi AF129742 MAT1-1-1::MAT1-2-1 HMG<br />

MAT1-1-1 α domain<br />

Cochliobolus cymbopogonis AF129744 MAT1-1-1 α domain<br />

AF129745 MAT1-2-1 HMG<br />

Sordariomycetes<br />

Gibberella zeae AF318048 MAT1-1-1 α domain<br />

MAT1-1-2 HPG domain<br />

MAT1-1-3 HMG<br />

MAT1-2-1 HMG<br />

Sordaria macrospora Y10616 Smt A-1 α domain<br />

Smt A-2 HPG domain<br />

Smt A-3 None identified<br />

Sm a-1 HMG<br />

Chaetomium globosum Broad Institute MAT1-1-1 α domain<br />

MAT1-1-2 HPG domain<br />

MAT1-1-3 HMG<br />

MAT1-2-1 HMG<br />

MAT1-1-2 like HPG domain<br />

Eurotiomycetes<br />

Emericella nidulans AY339600 MATB-1 (MAT-1, MATB) a α domain<br />

AF508279 MAT1-2 (MAT-2, MATA) a HMG<br />

a Names in parenthesis are different nomenclatures of the same gene<br />

gene, <strong>and</strong> between it <strong>and</strong> ORF1 (Fig. 15.2). Notably,<br />

thereisalsoagroupofself-compatibleisolates<br />

carrying only MAT1-1-1, a situation analogous to<br />

certain Neurospora self-compatible isolates (e.g., N.<br />

africana) whereonlythemat A-1 gene is detected<br />

(Glass et al. 1988, 1990). The remaining isolates of<br />

Stemphylium examined have undetermined sexual<br />

states <strong>and</strong> may be asexual, self-incompatible, or<br />

possibly debilitated selfers.<br />

These analyses indicate that mating-type<br />

gene arrangements are diverse in self-compatible<br />

Loculoascomycetes. A striking observation is that<br />

MAT1-2-1 was not detected in some Stemphylium<br />

isolates, which suggests that the HMG regulatory<br />

domain might be dispensable for sexual cycle<br />

control. Another possibility may be proposed,<br />

based on the recent finding that each MAT gene<br />

of Cochliobolus spp. appears to encode a protein<br />

with both HMG <strong>and</strong> α1activities(Lu<strong>and</strong>Turgeon,<br />

unpublished data; see Sect. III.C.1); the regulatory<br />

functions of the lost MAT1-2-1 gene may be taken<br />

on by the remaining MAT1-1-1 gene. Evolution<br />

of the target genes of MAT1-2-1 would then be<br />

necessary to bring them under the control of the<br />

HMG domain of MAT1-1-1. A first step for the<br />

validation of this hypothesis will be to demonstrate<br />

that Stemphylium MAT proteins have structures<br />

similar to those of Cochliobolus spp.<br />

c) Asexual Loculoascomycetes<br />

The evolutionary conservation of MAT genes in<br />

sexual Loculoascomycetes prompted Turgeon <strong>and</strong><br />

colleagues to ask if there were homologs of these<br />

genes in related species that had never been demonstrated<br />

to reproduce sexually (Sharon et al. 1996).<br />

This investigation was first conducted with Bipolaris<br />

sacchari, a pathogen of sugarcane. Among<br />

21 field isolates, 19 were demonstrated to carry a<br />

MAT1-2-1 gene <strong>and</strong> two carried a MAT1-1-1 gene.<br />

Recently Schoch, Saenz, <strong>and</strong> Turgeon (unpublished<br />

data) have updated this study, using more isolates<br />

<strong>and</strong> several molecular characters to investigate<br />

mating-type distribution <strong>and</strong> phylogenetic relationships<br />

of asexual B. sacchari. Combined data<br />

from both MAT genes, plus the 5 ′ MAT flank, the<br />

3 ′ MAT flank, <strong>and</strong> several additional sequences,<br />

confirm that B. sacchari isolates exist in the field<br />

as both mating types. All datasets indicated that<br />

the B. sacchari isolates fall into two well-supported<br />

groups, with representatives of both mating types<br />

in each group. The deduced MAT1-2-1 protein of

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