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Introduction to Fungi, Third Edition

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SACCHAROMYCES (SACCHAROMYCETACEAE)<br />

267<br />

Fig10.4 Saccharomyces cerevisiae.<br />

Diagrammatic summary of the processes of<br />

meiosis and ascospore delimitation (from<br />

Beckett et al.,1974). (a d) The spindle pole<br />

body replicates and the two new spindle pole<br />

bodies move <strong>to</strong> opposite poles of the nucleus.<br />

The nuclear membrane remains intact. (e,f)<br />

Further replication of the spindle pole bodies<br />

and rearrangement.The nuclear envelope is<br />

still intact. New membranes, the ascosporedelimiting<br />

membranes, form outside the spindle<br />

pole bodies. (g,h) Envelopment of the<br />

lobes of the dividing nucleus by the ascospore-delimiting<br />

membranes results in the<br />

formation of four haploid uninucleate<br />

ascospores.<br />

10.2.2 Mating in S. cerevisiae<br />

Many strains of S. cerevisiae are heterothallic,<br />

and the ascospores are of two mating types.<br />

Mating type specificity is controlled by a single<br />

genetic locus which exists in two allelic states,<br />

a and a, and segregation at the meiosis preceding<br />

ascospore formation results in two a and two a<br />

ascospores. Fusion normally occurs only between<br />

cells of opposite mating type, and this has<br />

been termed legitimate copulation. Such fusions<br />

result in diploid cells which can, under appropriate<br />

conditions, form asci with viable<br />

ascospores.<br />

The mating type (MAT) alleles are rather small<br />

and structurally similar <strong>to</strong> each other, but differ<br />

in their central region which comprises about<br />

650 base pairs (bp) in MATa and 750 bp in MATa<br />

(Fig. 10.5). Because of this difference, mating type<br />

alleles are often called idiomorphs. In haploid<br />

a-cells, the MATa locus expresses two genes, both<br />

of which encode regula<strong>to</strong>ry proteins. The a1 gene<br />

product interacts with a constitutively expressed<br />

protein not encoded by the MAT locus, Mcm1p,<br />

<strong>to</strong> activate several a-specific genes outside<br />

the MAT locus, notably those encoding the<br />

a-pheromone which is secreted by a-cells, and

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