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Brewer's Yeast: Genetics and Biotechnology

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Brewer's Yeast: Genetics and Biotechnology Applied Mycology and Biotechnology Volume 2. Agriculture and Food Production 1 @ 2002 Elsevier Science B.V. Al! rights reserved Brewer's Yeast: Genetics and Biotechnology Julio Polaina Instituto de Agroquímica y Tecnología de Alimentos, Consejo Superior de Investigaciones Científicas, Apartado de Correos 73, E46100-Burjasot (Valencia), Spain (E-Mail:jpola,[email protected]). The advance of Science in fue 19thcentury was a decisive force for fue deve10pmentand expansion of fue modero brewing industry. Correspondingly, fue brewing industry contributed important scientific achievements, such as Hansen's isolation of pure yeast cultures. Early studies on yeast were connected to fue development of different scientific disciplines such as Microbiology, Biochemistry and Genetics. An example of this connection is Winge's discovery of Mendelian inheritance in yeast. However, genetic studies with fue specific type of yeast used in brewing were hampered by fue complex constitution of this organismo The emergence of Molecular Biology allowed a precise characterization of fue brewer's yeast and fue manipulation of its properties, aimed at fue improvement of the brewing process and fue quality of fue beer. 1. INTRODUCTION The progress of chemistry, physiology and microbiology during the 19th.Century,allowed a scientific approach to brewing that caused a tremendousadvancement on fue production of beer. The precursor of such approach was the French microbiologist Louis Pasteur. At this time, fue Danish brewer Jacob Christian Jacobsen,algO founded fue Carlsberg Brewery and fue Carlsberg Laboratory. In Jacobsen'sown words, fue purpose ofthe Carlsberg Laboratory was: "By independent investigation to test the doctrines already furnished by Science and by continued studies to develop them into asfully scientij/c a basis as possible for the operation of mailing, brewing and fermentation". Louis Pasteur (1822-1895) demonstrated that alcoholic fermentation is a process caused by living yeast cells. His conclusion was that fermentation is a physiological phenomenon by which sugarsare converted in ethanol as a consequenceof yeast metabolismoIn 1876, Pasteur published "Etudes sur la Biere", which followed the trend of his previous book "Etudes sur le Vin", published ten years earlier. In Etudes sur la Biere, he dealt with fue diseasesofbeer and describedhow the fermenting yeast was often contaminated by bacteria, filamentous fungi, and other yeasts. However, the importance of Pasteurin relation with brewing is due to his discovery of yeast as fue agent of fermentation. His more specific contributions to this field are not to be considered among his greatest achievements.Probably, this had something to do with the fact that he did not like beer. Pasteur's work in connection with yeast and fue brewing industry has been recently reviewed by Anderson [1] and Bamett [2]. A crucial achievement for the developmentofthe brewing industry was accomplished by Emil Christian Hansen (1842-1909). Originally trained as a house painter and a primary school teacher, E. C. Hansen later became a botanist and a mycologist. In 1877, he was employed as a fermentation physiologist at fue Carlsberg Brewery. Familiar with fue work of Pasteur and facing fue problems of microbial contamination that often caused serious troubles in breweries, Hansen pursued the idea of 2 obtaining pille yeast cultures. To this end, he estimated the amount of yeast cells present in a beer sample. He made serial dilutions of fue sample until he reached an estimated concentration ofO.5 cells per mI, and used 1 mI aliquots afilie diluted suspensionto inoculate many individual flasks containing wort. After about a week oí incubation, roughIy half of fue cultures contained a single yeast colony, very few contained.two or more colonies, and no growth was observed in fue other half of fue flasks. Hansen concluded from this experiment that it was possible to obtain a single colony consisting of fue uncontaminated descendantsof . an individual cell. He performed additional experiments in which, starting with a mixture of two or more types of yeast, he was able to recover pille cultures of each different type. Another important contribution of Hansen to fue work with yeast was fue introduction of cultures on "solid medium". For this purpose he adapted fue procedure devised by Robert Koch for bacteria. Yeast colonies were grown on glass plates, on fue surface of a jellified medium prepared with gelatin. Hansen'snew techniques allowed him to obtain pille cultures of different brewing strains and also to characterizecontaminant strains that causeddifferent beer diseases.In 1883, fue Carlsberg Brewery started industrial production of lager beer with one of Hansen's pille cultures. This event became a milestone of the industrial revolution, since it meant fue transition from small-scale, artisan brewing to large-scale, modero production. The path led by the Carlsberg Brewery was soon followed by other companies, and in fue next few years fue technique of brewing with pille yeast cultures became standard in Europe and North America and caused an exponential growth of beer production all ayer the world. An exciting account ofthe work ofHansen has been given by van Wettstein [3]. 0jvind Winge was born in Arhus (Denmark) in 1886, shortly after the first industrial brewing with apure yeast culture. Winge was a very capable biologist who mastered different disciplines, including botany, plant and animal genetics, and mycology. In 1921, he became Professor of Genetics, firstly at the Veterinary and Agricultural University of Copenhagenand several years later at University of Copenhagen. Winge took the position of Director of fue Department of Physiology at fue Carlsberg Laboratory in 1933. When established in bis new position, he recovered fue collection of natural and industrial yeast strains gathered by Hansen and Albert Klocker, who both had preceded him at fue Department of Physiology. Winge faced fue problem that brewer's yeast strains were not able to sporulate, or did so very poorly, which made them unsuitable for genetic analysis. Therefore, he focused bis attention on baker's yeast (S. cerevisiae), which had long been a favorite organism for biochemical studies, and different varieties of Saccharomycescapable of sporulation (S. ludwigii, S. chevalieri, S. ellipsoideus, and others). With the help of a micromanipulation system of bis own design, Winge carried out dissection of fue asci of sporulated yeast cultures _andfollowed the germination of individual spores. He concluded that Saccharomyceshas a normal alternanceof unicellular haploid and diploid phases,i. e. it ... should behave genetically according to Mendel's laws. In collaboration with O. Laustsen, Winge reported the first results oftetrad analysis. After a lag period imposed by World War n, Winge starteda very productiveperiod that is markedby his collaborationwith Catherine o; Roberts. Together, they discovered fue gene that controls homothallism and many genesthat control maltose and sucrose fermentation. They also found that haploid yeast strains might have several copies afilie genesinvolved in fue fermentation ofthese sugars.They coined fue expression polymeric genes to designate a repeated set of genes that perform fue same function. The beginning of fission yeast (Schizosaccharomyces pombe) genetics is also linked to Winge. Urs Leupold spent a research stay in Winge's Department of Physiology where he established fue mating system and described the first cases of Mendelian inheritance for this yeast [4]. The work ofWinge in connection with yeast has been reviewed by R. K. Mortimer [5]. The birth of yeast genetics had a strong Scandinavian clout since besidesWinge, fue other prominent figure was Carl C. Lindegren, born in 1896 in Wisconsin, 3 USA, in a family of Swedish immigrants. The most transcendentachievement of Lindegren in connection with yeast genetics was fue discovery of fue mating types. This led to development of stable haploid cultures of both mating types and served to start fue cycle of mutant isolation and genetic crosses that made of Saccharomyces one of the most conspicuous organisms for genetic research. Other important achievements were fue discovery of fue phenomenonof gene conversion and the elaboration of fue first genetic maps of fue yeast. The work and the controversial personality of Lindegren have been the subject of an inspiring book chapter [6]. In 1847, the brewer J. C. Jacobsenstarted fue production ofbottom fermented (lager) beer at a brewery that he built in V alby, in fue outskirts of Copenhagen.He named bis brewery Carlsberg after bis five years old son Carl, who later became a maecenasof arts in Denmark, J. C. Jacobsenwas one of the pioneers of industrialization in Denmark. He introduced new procedures in fue brewing process that soon became standard and gave Carlsberg a rapid success.In 1875-76, J. C. Jacobsenestablished fue Carlsberg Foundation and fue Carlsberg Laboratory. The Carlsberg Laboratory was divided in two Departments, Physiology and Chemistry. As a tradition, both Departments have focused their work main1y, albeit not exclusively, on processesand organisms of special significance for brewing, s'uch as yeast and barley. The first director of fue Department of Chemistry was Johan Kjeldah1, who invented fue procedure for fue determination of organic nitrogen that carries bis llame. Undoubtedly, fue most popular contribution of the Department of Chemistry was fue concept of pH, due to S0ren P.
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