In this review we describe the role of the yeast Saccharomyces in the development of human societies including the use of this organism in the making of wine, bread, beer, and distilled beverages. We also discuss the tremendous diversity of yeast found in natural (i.e., noninoculated) wine fermentations and the scientific uses of yeast over the past 60 years. In conclusion, we present ideas on the model of "genome renewal" and the use of this model to explain the mode by which yeast has evolved and how diversity can be generated.
Genetic and Molecular Studies of the Yeast GenomeIn this discussion the term yeast is used rather loosely because there are hundreds of species of yeast. However, we refer to only a few species in this article, Saccharomyces cerevisiae, Saccharomyces carlsbergensis, Saccharomyces bayanus, and Saccharomyces paradoxus, which are used in the making of wine, bread, beer, and distilled beverages. Taxonomically, yeast is defined as a single-cell eukaryotic microorganism that usually divides by budding.Although humans have used yeast for centuries (see Box 1), the yeast S. cerevisiae has only been studied at the genetic level since the mid 1930s with the pioneering work of Winge (1935). He used as his starting material, strains of Saccharomyces that had been isolated many years before by Hansen (1891). Lindegren, also one of the pioneers of yeast genetics, used a strain, EM93, that had been isolated by Emil Mrak (Mortimer and Johnston 1986). This strain was found on rotting figs in Merced, California, in 1938, and turned out to be heterothallic (i.e., having two types of sterile haploid individuals which are fertile when together). Other strains found on the same figs were homothallic (one haploid phase that produces gametes capable of fusing to create a zygote).The strains that Winge studied were nearly all homothallic, which required particular skills in the making of crosses. Lindegren was fortunate in being given a heterothallic strain, constructing crosses was very easy, and eventually this led to the development of modern yeast genetics. However, most of the studies of yeast genetics and, subsequently, yeast molecular biology, were carried out on a very few strains, most of which were derived from Lindegren's original strain EM93. Strain S288C, used to sequence the yeast genome, was constructed in the early 1950s (Mortimer et al. 1957;Mortimer and Johnston 1986). It has been estimated that Ն85% of the genome of S288C comes from EM93 (Mortimer and Johnston 1986). It seems likely that this original diploid strain was also a wine yeast (∼10% of wine yeast are heterothallic) and was carried to the rotting figs by insects; the yeast S. cerevisae does not travel independently through the air.Genetic research on the brewing yeasts shows that most have extremely low spore viabilities. Other research suggests that some of them may be polyploid or aneuploid (Johnston 1990). These characteristics would contribute to strain stability and this property may have been inadvertently selected by the brew...