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Introduction to Fungi, Third Edition

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SACCHAROMYCES (SACCHAROMYCETACEAE)<br />

265<br />

the enormous importance of S. cerevisiae for<br />

fundamental cell biology. This fungus was the<br />

first eukaryote (in 1996) <strong>to</strong> have its complete<br />

genome sequenced, and this <strong>to</strong>gether with the<br />

ease of genetic manipulation has further<br />

enhanced the status of S. cerevisiae as a workhorse,<br />

if not ‘model organism’, for eukaryote<br />

research.<br />

10.2.1 The life cycle of S. cerevisiae<br />

Vegetative cells of S. cerevisiae are generally<br />

diploid in nature, although tetraploid or aneuploid<br />

strains also occur. Strains may be homoor<br />

heterothallic. The chromosome number is 16<br />

(Cherry et al., 1997). The life cycle of S. cerevisiae<br />

is presented in Figs. 10.2 and 10.3. The haploid<br />

ascospores often fuse within the ascus where<br />

they were formed (Fig. 10.3d), or shortly after<br />

release. However, if individual ascospores<br />

become isolated, they can germinate and reproduce<br />

as haploid cells by budding. Where two<br />

haploid cells of opposite mating type are in close<br />

contact, they secrete peptide hormones and produce<br />

plasma membrane-bound recep<strong>to</strong>rs which<br />

recognize the hormone of opposite mating type.<br />

The binding of a hormone molecule <strong>to</strong> the<br />

matching recep<strong>to</strong>r sets a signalling chain in<br />

motion (reviewed by Bardwell, 2004) which<br />

arrests the mi<strong>to</strong>tic cell cycle at G1, stimulates<br />

transcription of mating-specific genes and<br />

causes polarized growth of the two cells <strong>to</strong>wards<br />

each other (Leberer et al., 1997). Mating initially<br />

involves an increased ability of the surfaces of<br />

two cells <strong>to</strong> adhere <strong>to</strong> each other. This so-called<br />

sexual agglutination is mediated by glycoproteins.<br />

It seems that these are components of<br />

fimbriae, i.e. long filaments radiating outwards<br />

from the cell wall (see Fig. 23.15). Agglutination<br />

is followed by co-ordinated digestion of the<br />

walls separating the two cells. Plasmogamy and<br />

karyogamy follow swiftly (Gammie et al., 1998).<br />

The resulting diploid cell (Fig. 10.3e) can carry on<br />

reproducing asexually by budding. In contrast<br />

<strong>to</strong> Schizosaccharomyces pombe, there are therefore<br />

two mi<strong>to</strong>tic cycles in the life cycle of S. cerevisiae.<br />

Under optimum conditions, the culture doubling<br />

time by mi<strong>to</strong>sis is about 100 min.<br />

Diploid strains of S. cerevisiae can be induced<br />

<strong>to</strong> form ascospores by suitable treatment, and<br />

this yeast is therefore termed an ascosporogenous<br />

yeast, in contrast <strong>to</strong> asporogenous yeasts<br />

in which ascospores have not been observed.<br />

Meiosis can be induced by growing the yeast on<br />

a nutrient-rich presporulation medium containing<br />

an assimilable sugar, a suitable nitrogen<br />

source for good growth (nitrate is not utilized),<br />

Fig10.2 The life cycle of S. cerevisiae. Both haploid and diploid cells can reproduce by budding.Open and closed circles represent<br />

haploid nuclei of opposite mating type; diploid nuclei are larger and half-filled. Key events in the life cycle are plasmogamy (P),<br />

karyogamy (K) and meiosis (M).

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