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Yeast Biotechnology: Diversity and

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Yeast Biotechnology: Diversity and Applications
Mustafa Trker
R&D, Environment and Quality Coordination Manager
Pakmaya, Kocaeli, Turkey
The yeasts have been exploited by mankind for thousands of years for food and fermentation processes.
Traditionally the yeast has been used for the production of alcoholic beverages, biomass and glycerol.
Saccharomyces cerevisiae has been described as mankinds most domesticated organism and still widely
exploited yeast species in industry today. The number of yeast species described so far is about 1500 and
only about a dozen is used at industrial scale. Some 70-80 species have been shown to possess potential
value for biotechnology. According to modest estimate, known yeast species represent roughly 5% of the
total number that may inhabitat Earth surface. Modern applications of yeasts have been greatly expanded
beyond clasical applications. Yeasts, especially S. cerevisiae and other non-saccharomyces yeasts today are
increasingly used for the heterologous production of enzymes and pharmaceutical proteins. Yeasts have
important roles in environmental applications such as bioremediation and removal of heavy metals from
wastewaters. Yeasts are also used in agriculture as biocontrol agents. Several chemicals can be produced
using yeast as a biocatalyst. New developments in engineering yeast have introduced novel capabilities to
extend substrate range and produce new products so far yeast can not produce. S. cerevisiae is largest
cultivated organism so far. Having in mind diversity and potential of all yeast species, the cultivation and
utilization of Saccharomyces yeasts are still the tip of the iceberg and there is a vast potential yet to be
discovered for the production of valuable products using saccharomyces and non-saccharomyces yeasts.
Introduction
Yeasts have been used for making bread, beer and wine since ancient ages. First
microscopic observation of microbial cells and description of the microscobic appearance of
yeast were observed by A. van Leeuwenhoek in 1680. Their role in fermentation was
recognized by Pasteur, and the first pure cultures (starters) of brewers and wine yeast were
obtained by Hansen and Mller-Thurgau, respectively, at the end of the 19th century. Since
then the application of yeast starters has become a standard practice in the industrial
fermentation not only for food and beverages but also for a broad variety of other products
made by yeasts or from yeast cells. The traditional fermentation processes are carried out by
a single species of yeasts, Saccharomyces cerevisiae , hence for many, its name is
synonymous with yeasts, and it is thought that all yeasts are fermentative. Contrary to
general belief, about half of described species not being able to ferment; nevertheless many
of these have gained a significant role in biotechnology (Barnett and Barnett, 2011).
Yeasts form an artificial group of fungi comprising mostly unicellular organisms reproducing
vegetatively by budding and can be classified either to Ascomycetes (e.g. Saccharomyces,
Candida ) or Basidiomycetes (e.g. Filobasidiella, Rhodotorula). About 74,000 species of fungi
including yeasts have been described, while estimates of the total number of fungal species
has been estimated to be as high as 1,500,000 (Hawksworth 2004), Thus, fungi are among
the richest kingdoms on earth with respect to biodiversity, but much work is needed to
understand their potential to provide valuable industrial resources. Currently, there are
approximately 1,500 recognized yeast species listed in the latest edition of The Yeasts: a
Taxanomic Study (2011) and estimated total number is around 150,000 forming roughly 5-
10% of estimated fungal species. In the previous edition of this book, in 1998, 700 species
were described. Since then, the number of recognized yeast species has doubled, with a
27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Advances in Science and Industrial Productionss of Bakers Yeast
Page 1
steep increase particularly in the number of the basidiomycetous yeasts. Of all these yeast
species, only about a dozen is used at industrial scale, and some 70 80 species have been
shown at laboratory scale to possess potential value in biotechnology (Kurtzman et.al.,
2011; Deak, 2009).
The most important nutrients for yeasts are carbohydrates that serve for both carbon and
energy sources. Only a few sugars, mostly hexoses and oligosaccharides, can be fermented
by yeasts. The range of carbon sources that can be utilized aerobically is much wider, and
includes hexoses, pentoses, alcohols, organic acids, and other carbon compounds. Mono-
and oligosaccharides are widely utilized by yeasts, although the fermentation of galactose is
limited to some species. Not all yeasts are able to metabolize certain di- and trisaccharides
(sucrose, maltose, lactose, or raffinose) for lack of the necessary hydrolytic enzymes.
Utilization of pentoses is also restricted among yeasts. The fermentation of xylose may be a
potentially useful property for the industrial production of ethanol from hydrolysis products
of plant hemicelluloses. The ability of yeasts to metabolize polysaccharides and complex
carbohydrates is restricted to relatively few species. Utilization of starch is of particular
interest for industrial production of yeast biomass (single-cell protein, SCP) from starchy
agricultural wastes. Db. occidentalis and Lipomyces species are among the few types of
yeasts possessing enzymes of various amylase activities. S. cerevisiae is unable to hydrolyze
starch. Other possible carbon sources for yeasts are hydrocarbons, and several species are
capable of growing on these compounds (Deak, 2008). Both organic and inorganic nitrogen
sources can be utilized by yeasts. Although very few species can hydrolyze proteins
extracellularly, short peptides can be transported into the cell and utilized intracellularly.
Amino acids, amines, and urea are suitable nitrogen sources for practically all yeasts (Large,
1986), in addition to inorganic ammonium salts. Nitrate utilization, however, is confined to
certain species or genera of yeasts, and this is a valuable diagnostic character used for
identification purposes (Rossi and Berardi, 2009).
Yeasts are usually tolerant to broad pH ranges and are often found at the acidic end of the
scale where many bacteria are not able to compete. In terms of maximum temperature,
Tmax, microorganisms can be subdivided into three groups. The term thermophilic is applied
to microorganisms whose Tmax is well above 50C; those capable of growth at Tmax between
25C and 50C are referred to as mesophilic; and psychrophiles are those classified as
having a Tmax below 25C. Considered in these terms, nearly all known yeasts are mesophilic,
and grow best between 20C and 30C. However, truly thermophilic yeasts remain unknown.
Rarely can they grow above 42 0C and those that do may occupy specialized habitats such as
the intestine of warm-blooded animals. Most strains of S. cerevisiae occurring widely in
industrial fermentation can grow at 37C, whereas growth in a similar environment of S.
bayanus is limited up to 3035C. The range of yeasts able to grow above 40C is limited.
Most of isolated thermotolerant yeasts capable of growing at temperatures above 40C
belonged to Kluyveromyces marxianus, but other thermotolerant strains were identified
with P. polymorpha, Geo. capitatum, S. cerevisiae, and Candida and Debaryomyces species.
Kluyveromyces strains are noteworthy for their relatively high Tmax values. However,
temperatures above 50C are usually lethal for yeast cells (Deak, 2008; Starmer and
Lachance, 2011).
Role of Yeast in Traditional Food Fermentations

Page 2
Traditionally the yeast has been used for the production of alcoholic beverages, biomass and
glycerol. Saccharomyces cerevisiae has been described as mankinds most domesticated organism
and still widely exploited yeast species in industry today. Although the production of alcoholic
beverages and leavening of dough have been practiced by several millenia, glycerol was
produced by S. cerevisiae by chemical diversion of central metabolic pathways and the
process was introduced before the second world war.
Alcohol
Yeasts
Glycerol Food
Figure-1: Traditional Uses of Yeasts (adapted from Walker, 1998)
A fermented food or beverage is defined as an edible product prepared from the raw or
cooked materials of plant or animal origins by microorganisms either naturally or by adding
mixed or pure culture(s). The essential objective of food fermentation is to carry over
supplies from the time of plenty to those of deficit. Traditionally people knew how to culture
the beneficial microorganisms, mostly lactic acid bacteria, yeasts and filamentous moulds,
for production of foods for consumption. What was the scientific explanation and identity of
these microorganisms were unknown to them. Microorganisms are present in or on the
ingredients, utensils, environment, and are selected through adaptation to the substrate for
fermentation. Rice-soybean-fish-alcoholic beverage diet is the characteristic food culture
of the East and South East Asia, whereas wheat-milk and milk products-meat-wine is the
basic diet in the Western part of the world. In East Asia rice is a staple food whereas in West
Asia wheat or barley is a staple food. Milk products similar to West and alcoholic beverages
similar to East are encountered in food habits of the people of the Himalayas. In Asia,
moulds are predominant microorganisms in the fermentation processes, whereas in Africa,
Europe and America, fermented foods are prepared exclusively using bacteria or bacteria-
yeasts mixed cultures; moulds seem to be little or never used. However, in the Indian sub-
continent, mostly due to wide variation in agroclimatic conditions and diverse form of
dietary culture, all major groups of microorganism (bacteria-yeasts-moulds) are associated
with fermented foods showing the transition of food culture (Tamang and Fleet, 2009; Fleet,
2011).
Yeasts play vital roles in production of many traditional fermented foods and beverages
across the world signifying the food culture of the regions and the community. Functional
yeasts genera associated with fermented foods and beverages are mostly Brettanomyces (its
Page 3
perfect stage, Dekkera ) , Candida, Cryptococcus, Debaryomyces, Galactomyces, Geotrichum,
Hansenula, Hanseniaspora (its asexual counterpart Kloeckera ), Hyphopichia, Kluyveromyces,
Metschnikowia, Pichia, Rhodotorula, Saccharomyces, Saccharomycodes, Saccharomycopsis,
Schizosaccharomyces, Torulopsis, Trichosporon, Yarrowia and Zygosaccharomyces. The
common fermented foods and beverages, mostly prepared by yeasts, or in combination with
bacteria and moulds are: beers and ales, breads and bakery products, cachaa, cheeses,
kenkey, kimi, dairy products (e.g., kefir; yoghurt, fermented milk), cocoa, coffee, fermented
meat and sausages, fermented olives and cucumber, soy paste, tea fungus, silage and
probiotics.
Many traditional foods and beverages in Anatolia are produced by fermentation involving
either bacteria including lactic acid bacteria or yeasts or mixtures of both. Lactic acid
fermentation in particular plays an important and predominant role in the production of
traditional foods and beverages of Turkey. Tarhana, kefir, koumiss and boza are traditional
fermented foods in which yeasts are present together with lactic acid bacteria. The yeast
species in these traditional fermented foods are Kluyveromyces marxianus, Candida
inconspicua, Candida maris, Torulopsis kefir Saccharomyces cerevisiae, Saccharomyces lactis, Torula
koumiss Kluyveromyces lactis, non-lactose fermenting yeast such as Saccharomyces unisporus,
Saccharomyces uvarum, Candida tropicalis, Candida glabrata, Geotrichium penicillatum, and
Geotrichium candidum, Candida diverca, Candida inconspicua, Candida pararugosa, Issatchenkia
orientalis (Kabak, B. And Dobson, 2011).
Food and Feed Yeasts

Yeasts have long been cultivated as rich sources of protein, minerals, vitamins (particularly B
vitamins), and other nutrients for humans and animals. Several yeast species have been used
for biomass production, including C. utilis, other non-methylotrophic Candida spp.,
Saccharomycopsis (Endomycopsis) fibuligera, Kluyveromyces spp., and S. cerevisiae.
Methylotrophic, ethanol-utilizing, and fat- and hydrocarbon-utilizing yeasts including species
of Candida, Ogataea, Pichia, and Trichosporon have also been used for biomass production.
Production of yeast SCP has certain advantages compared to plant, animal, and other
microbial sources of SCP including rapid growth and accumulation of biomass, high protein
content (up to 50%), high contents of vitamins and minerals, and ability to grow on a wide
variety of substrates, including various industrial waste streams. The production of value-
added SCP from inexpensive substrates could help to alleviate world shortages in the food
supply, particularly in developing countries. Substrates utilized have included molasses,
starch, cassava, Jerusalem artichoke, whey products, sulfite waste liquor, potato wastes,
brewery wastes, and other waste streams from agricultural processes, food processing, and
industrial processes. Due to the relatively large cell size and flocculation abilities of yeasts,
they can be more easily harvested than bacteria from the fermentation liquor. Compared to
bacteria, many yeasts contain low quantities of nucleic acids, which can have detrimental
nutritional effects. The nutritional value and safety of SCP has received some evaluation
(Boze et.al., 1992; Bekatorou, et.al, 2006).
Certain yeast species have been used as prebiotic and probiotic agents for preventing or
treating various intestinal, nutritional, and toxicological disorders. In particular,

Page 4
Saccharomyces boulardii, originally isolated from fruit in Indochina, has been used for
treatment of intestinal diseases in children and adults since 1950. Recent studies have
shown that it is conspecific with S. cerevisiae, but S. boulardii appears to have certain
genotypic and phenotypic properties that may contribute to its probiotic properties.
Controlled clinical trials have shown efficacy of S. boulardii for prevention or elimination of
several intestinal disorders, including infection by Clostridium difficile associated with
antibiotic-induced diarrhea. Yeasts have also been hypothesized to prevent other intestinal
infections, including inflammatory bowel disease and Crohn's disease. The mechanisms of
probiotic activity have not been elucidated but may involve alteration of inflammatory and
immune responses or destruction of toxic factors. Although yeasts may be promising
therapeutic agents, they need to undergo controlled clinical trials to critically evaluate their
efficacy. Other yeasts allowed and commonly used in animal feeds as probiotic additives are
Candida pintolopesii, C. saitoana and S. cerevisiae(Bekatorou, et.al, 2006). Preliminary
evidence indicates that certain yeasts can produce prebiotics, which are compounds
(generally oligosaccharides) that stimulate the growth of bifidobacteria and other beneficial
bacteria in the gut of humans and animals or oligopeptides that have beneficial health
benefits by stimulation of immune response. Prebiotics are often sugar derivatives such as
fructooligosaccharides, of which certain yeasts can catabolize due to their formation of
inulinases. K. lactis was found to synthesize prebiotic oligosaccharides and
immunostimulatory peptides from whey (Belem and Lee 1998; Bekatorou, et.al, 2006;
Torres et.al., 2010).
Yeast glucans and cell wall polysaccharides have been used as adjuncts for animal and fish
feeds. These polysaccharides have been shown to promote animal growth and health by
various mechanisms including immunomodulation, oxidative status, binding of toxins and
pathogens, and interactions with gut constituents (Kim et.al, 2011). Glucans and mannans
have also been found to have a myriad of biological functions in animal models and
potentially in humans, including modulation of histamine release and antitumor activities.
Further studies are needed in this area to evaluate health promotion by yeast cell wall
polysaccharides (Zechner-Krpan et.al., 2009).
Modern Applications of Yeasts

The annual world production of S. cerevisiae is at a level that exceeds the combined
production of all other industrial microorganisms by about two orders of magnitude. The
economic values of fermented beverages and foods involving yeasts are enormous. Although
the yeast is synonymous with Saccharomyces cerevisia, the other yeast species of
biotechnological importance have been introduced to produce industrial products beyond
traditional foods as shown in Table 1. They have wide-ranging fundamental and industrial
importance in scientific, food, medical, and agricultural disciplines (Fig. 3.1). In addition to
traditional industrial applications of yeasts in several food fermentations such as alcoholic
beverages, bakery products, cheese, sausages, and other fermented foods. modern
applications of yeasts involve the production of fuel ethanol, single cell protein (SCP), feeds
and fodder, industrial enzymes, and small molecular weight metabolites. More recently,
Komagataella (Pichia) pastoris, Saccharomyces cerevisiae, Ogataea (Hansenula) polymorpha,
and certain other yeast species have been developed as industrial organisms for the
heterologous production of enzymes and proteins, including protein pharmaceuticals.
Yeasts, especially S. cerevisiae, are increasingly being used as hosts for expression of protein

Page 5
biocatalysts and multi-enzyme pathways for the synthesis of fine chemicals and small
molecular weight compounds of medicinal and nutritional importance. Yeasts have
important roles in agriculture as agents of biocontrol, bioremediation, and as indicators of
environmental quality. Several of these processes and products have reached commercial
utility, while others are in development (Branduardi and Porro, 2012; Johnson, 2013a,b;
Johnson & Echavarri-Erasun, 2011).
Table 1: Biotechnologically Important Yeast sp. (Johnson, 2013a,b)

Ascomycetous Basidiomycetous
Saccharomyces cerevisiae Rhodotorula spp.
Schizosaccharomyces pombe Rhodosporidium spp.
Kluyveromyces lactis Trichosporon spp.
Kluyveromyces marxianus Xanthophyllomyces dendrorhous
Schwanniomyces occidentalis Cryptococcus spp.
Lipomyces spp. Phaffia rhodozyma
Saccharomycopsis spp.
Debaryomyces hansenii
Ogataea polymorpha
Komagataella pastoris
Scheffersomyces stipitis
Pichia spp.
Yarrowia lipolytica
Candida spp.
Blastobotrys adeninivorans

Page 6
TRADITIONAL YEAST
FERMENTATIONS
Beer, Wine, Sake, Soy Sauce,
Other Food Fermentations FOOD AND FEED
ENVIRONMENTAL
BIOTECHNOLOGY INGREDIENTS
Bioremediation, Pollutant Enzymes, Flavors, Pigments,
Degradation Amino acids, Organic acids
BIOCONTROL BIOFUELS
Crop protection, Food Bioethanol, Biodiesel, FAEE,
and Feed Safety, Biobutanol, sesquiterpenoids
Probiotics Yeast farnesene, bisabolene, and
amorphadiene
Biotechnology
BIOMEDICAL RESEARCH
Drug discovery, Drug BIOCATALYSIS
Resistance and Pharmaceuticals, Chiral
Metabolism, Elucidation Chemical Intemediates,
of Disaese Mechanism Biotransformations
FUNDAMENTAL BIOLOGICAL
RESEARCH HETEROLOGOUS PROTEIN
Molecular and Cellular Biology, PRODUCTION
Genomics, Functional Genomics, Protein Pharmaceuticals,
Pathway Engineering, System Biology Enzymes, Hormones,
Mechanisms Vaccines, Toxins
Figure-2: Modern Applications of Yeasts (adapted from Walker, 1998)
Food and Feed Enzymes from Yeasts

The term enzyme (literally in yeast) was coined by Khne in 1876. In 1907, Eduard
Bchner received the Nobel Prize in Chemistry for his discovery of cell-free fermentation
using bakers yeast cell-free extracts. Enzyme technology utilizing cell-free enzymes and
whole-cell biocatalysts is integral to several large-scale chemical, pharmaceutical, and
agricultural bioprocesses (Aehle 2007). The 2010 market estimate for technical and food and
feed enzymes was around USD 3.5 billion (52% technical enzymes and 48% food and feed
enzymes), leaving a figure of around USD 2 billion for other enzymes. It has been forecasted
to increase to 7 billion by 2013 and to 8 billion by 2015, representing an average growth rate
of 6.8% (Illanes et.al., 2014). The enzyme market is generally categorized into four utility
classes: (a) technical enzymes, which comprise about 65% of the market, including enzymes
used in the detergent, starch, textile, leather, pulp and paper, and personal care industries;
(b) food enzymes, about 25% of the market, including enzymes used in brewing, dairy, wine
and juice, fats and oils, and baking industries; (c) feed enzymes, about 10% of the market,
used in animal feeds; and (d) diagnostic enzymes, comprising a small portion of the overall
enzyme market. In recent years, growth of the bulk enzyme industry has been particularly
robust in the baking and animal feed sectors, in industrial organic syntheses for fine
chemicals and pharmaceuticals, and to a lesser degree for paper and pulp processing,
production of biofuels, and for personal care (Aehle 2007). The food industry is the largest

Page 7
user of bulk enzymes, accounting for about 45% of total use. The bulk enzyme market is
dominated by enzymes produced by bacteria and filamentous fungi cultivated in submerged
culture, particularly Bacillus and Aspergillus species (Aehle 2007). Only a limited number of
yeast enzymes are currently used as bulk enzymes in commodity processes, as shown in
Table 2 (Johnson & Echavarri-Erasun, 2011).
Table 2: Industrial Enzymes from Yeasts (Johnson & Echavarri-Erasun, 2011)

Enzyme Yeast Industry
Chymosin Kluyveromyces spp. Food processing
Saccharomyces cerevisiae
-Galactosidase Saccharomyces spp Feed applications
L-Glutaminase Zygosaccharomyces rouxii Therapeutic
Analytical
Inulinases Candida spp. Food applications
Kluyveromyces marxianus
Invertase Saccharomyces cerevisiae Food applications
Lactase Candida pseudotropicalis Food processing
Kluyveromyces spp.
Lipase Candida rugosa Food processing
Pseudozyma antarctica A, B Flavors, Wastewater
Geotrichum candidum Degreasing, Bioremediation
Trichosporon fermentum Therapeutic, Detergent
Yarrowia lipolytica
L-Phenylalanine Rhodotorula spp. Pharmaceutical
ammonialyase Rhodosporidium spp.
Phenylalanine Candida boidinii Pharmaceutical
dehydrogenase
Phytase Ogataea polymorpha Feed
Nutrition
However, several yeast enzymes have found application in the production of high-value
specialized fine chemicals and for biotransformation of pharmaceutical intermediates (see
section 13). Currently, about 90% of industrial enzymes are produced as heterologous
proteins by recombinant methods. Examples of industrially-relevant recombinant enzymes
produced in yeasts are presented in Table 3.
Table 3: Examples of Industrial Recombinant Enzymes

Produced in Yeasts (Johnson & Echavarri-Erasun 2011)
Enzyme Recombinant Yeast Host
Chymosin Kluyveromyces lactis
Glycolate oxidase Komagataella pastoris
Phytase Komagataella pastoris
Certain yeasts, especially Komagataella pastoris and S. cerevisiae, are increasingly being
utilized for heterologous production of enzymes and a variety of other proteins (Aehle 2007,
van Beilen and Li, 2002). Bla. adeninivorans, O. polymorpha, K. lactis, Schiz. pombe, Y.
lipolytica, Pseudozyma spp., and other yeast species have also been developed for the
production of heterologous proteins (Demain and Vaishnav, 2009; elik and alk, 2012).
Industrial recombinant enzymes and those close to commercialization produced by yeasts
are presented in Table 2 and 3. Yeasts are desirable hosts of enzymes for food uses because
of their lack of production of toxic secondary metabolites (Olempska-Beer et al. 2006). K.

Page 8
lactis is currently approved by the FDA for production of recombinant enyzmes (chymosin)
for uses in foods. Whole cells used as biocatalysts for high-value pharmaceuticals and fine
chemicals is a rapidly growing sector of the enzyme industry.
Yeast Metabolites and Bulk and Fine Chemicals

Several yeast species produce commodity chemicals in levels at or approaching commerical
utility. The uses of yeasts in White Biotechnology applications, or production of low-cost,
high-volume products is accelerating. S. cerevisiae, K. lactis, and Schef. stipitis have been
developed for the production of L-lactic acid. The availability of lactic acid from yeasts and
other organisms has led to expanded uses such as its polymerization to polylactic acid.
Polylactic acid and related polymers are under development for a variety of products
including biodegradable plastics and textile fibers (Branduardi et.al, 2008; Branduardi and
Porro, 2012). Saccharomyces cerevisiae and Rh. glutinis have been used for medium-scale
production of amino acids including lysine, methionine, phenylalanine, and proline. Yeasts
have also been used for the production of alcohols used in bulk processes. C. magnoliae, C.
peltata, and a Candida sp. produce erythritol in 41 to 56% yields from glucose or xylose in an
approximately 3-day fermentation. Mannitol, widely used in the food, pharmaceutical and
chemical industries, is produced by various yeast species from glycerol. Glycerol is a
commodity chemical that is used in a number of industrial products. It has been traditionally
produced by chemical synthesis from propylene, and is also recovered following hydrolysis
of fats. C. glycerinogenes produces glycerol (144 g/l) from glucose in a 72-hour batch
fermentation (Johnson & Echavarri-Erasun 2011). S. cerevisiae produces glycerol during
ethanol formation, and synthesis is increased by osmotic stress (Attfield et.al., 2003). In the
early 1900s, Carl Neuberg found that glycerol production in S. cerevisiae was enhanced by
chemical diversion of central metabolic pathways. To increase the cellular availability of
more reducing equivalents (NADH/NADPH) for glycerol production, agents such as sulfite or
alkali were added to the fermentations to inhibit the conversion of acetaldehyde to ethanol,
and to shuttle carbon flow to glycerol. The use of sulfites may have utility in other
fermentations, such as in the fermentaion of xylose, in which availability of reducing
equivalents appears to limit production of high yields of ethanol (Jeffries, 2006) and in
astaxanthin formation by X. dendrorhous in which biosynthesis requires high quantities of
reducing equivalents. Yarrowia lipolytica has been used for the industrial production of
organic acids including -ketoglutaric, pyruvic, citric and isocitric acids, which are used in
food products (Sauer et.al., 2008; Liu et.al., 2013). Citric acid is a high-volume commodity
fermentation product, second only to ethanol (Levinson et al. 2007). The ability of Y.
lipolytica and certain Candida spp. to form high quantities of organic acids from various
substrates, such as sucrose and glycerol, has seen renewed interest. Other yeast species,
including C. glabrata, have been used industrially for the production of pyruvic acid. Several
yeasts have attracted industrial interest for production of primary chemicals such as itaconic,
malic, gluconic, brassylic, sebacic, and fumaric acids (Walker 1998), but several of the
processes are presently non-competitive with other commercial chemical or fermentation
processes. Yeasts have been used for production of vitamins. Pichia guilliermondii
overproduces riboflavin from xylose and ribitol in response to iron deprivation in the
medium (Sibirny and Boretsky, 2009). Pichia sp. have been investigated for the production
of pyridoxine (Walker 1998). Several yeast species produce glycolipids and surfactants that
could have applications in the oil, food, cosmetic, and pharmaceutical industries (Amaral,

Page 9
2008). The representative list of commodity chemicals produced by yeast species are shown
in Tbale 4.
Table 4: Comodity Chemicals Produced by Yeasts (Amaral, 2008; Branduardi and Porro,
2012; Sauer et.al., 2008)
Products Yeast
Amino acids including lysine, methionine, Saccharomyces cerevisiae and Rh. glutinis
phenylalanine, and proline
Erythritol C. magnoliae, Moniella spp.
C. peltata, and a Candida sp.
Mannitol Various yeast sp.
Glycerol C. glycerinogenes
S. cerevisiae
Astaxanthin Xanthophyllomyces dendrorhous, Phaffia
rhodozyma
-ketoglutaric, pyruvic, citric and isocitric acids, Yarrowia lipolytica
Gluconic acid Aureobasidum pullulans
2-phenylethanol Pichia fermentans
Pyruvic acid Candida glabrata
Riboflavin Pichia guilliermondii
Glycolipids and surfactants, Sophorolipids Basidiomycetous yeasts, Pseudozyma,
Candida, Kurtzmanomyces
easts are well-recognized as sources of food flavor enhancers for foods including nucleotides
and yeast extracts and autolysates, ribonucleotides, cell wall mannoproteins, cell wall
glucans, edible proteins, choline, glycerol, and inositol, mineral-enriched yeast, sterols,
vitamins, whole cell or single-cell proteins or single-cell oils, yeast enzymes (Walker, 1998).
Nucleotide taste enhancers are also produced by the enzymatic hydrolysis of yeast RNA into
nucleotide phosphates. Yeast cell wall polysaccharides have been used as adjuncts for
animal and fish feeds. These polysaccharides have been shown to promote animal growth
and health by various mechanisms including immunomodulation, oxidative status, binding of
toxins and pathogens, and interactions with gut constituents (Abbas, 2006). While chemical
synthesis is currently the preferred technology for producing flavor compounds, increasing
demand for natural flavors has given impetus to the development of microbial systems for
the production of VOSCs (volatile organic sulfur compounds) by ascomycetous and
basidiomycetous yeasts (Buzzini et.al, 2005).
Biocatalysis and Fine Chemicals

Catalysis in the chemical and pharmaceutical industries has traditionally been dominated by
chemical syntheses using non-biological catalysts. However, several disadvantages are
inherent to certain industrial chemical synthetic processes, including relatively low catalytic
efficiency for many reactions, lack of enantiomeric specificity for chiral syntheses, the need
for reaction conditions of high temperature, low pH and high pressure requirements, as well
as the extensive use of organic solvents with consequent formation of organic waste and
pollutants. These concerns and drawbacks have stimulated keen interest in biocatalytic
processes using enzymes. Advantages of enzymes for chemical processes include mild
reaction conditions, i.e., operation at temperatures from 20 to 80 0C, neutral or slightly
acidic pHs, and atmospheric conditions.

Page 10
Although a large number of studies and a voluminous literature have been devoted to the
production of fine chemicals using enantioselective catalysis by enzymes, relatively few
enzymatic processes are currently used on an industrial scale. It has been estimated that
about 150 industrial syntheses currently utilize enzymes or microbial whole cell catalysts
(Straathof et al. 2002). Several yeasts produce enzymes and bioacatalysts of realized and
potential value in the traditional enzyme industry and as chiral-specific biocatalysts for the
fine chemical and pharmaceutical industries (Aehle 2004, Liese et al. 2006). Lipases and
esterases, commonly produced by yeasts, are the most frequently used biocatalysts in
industrial organic syntheses (Kapoor and Gupta, 2012; Brgidaa et.al., 2014; Wache et.al.,
2006; Fickers et.al., 2011). Table 5 shows examples of commercially used industrial
biotransformations employing wild type yeast whole-cell biocatalysts.
Table 5: Industrial biotransformations employing wild type yeast whole-cell biocatalysts

(adapted from Pscheidt & Glieder, 2008)
Yeast strain Enzyme name Product Company
Zygosaccharomyces Alcohol NAD+ Benzodiazepine Eli Lilly and
rouxii oxidoreductase Company, USA
Geotrichum Dehydrogenase, chiral -hydroxy ester Bristol-Myers Squibb,
candidum NADPH-dependent USA
Candida sorbophila Dehydrogenase (R)-amino alcohol Merck & Co, Inc.,
USA
Pichia methanolica Reductase Ethyl-5-(S)-
hydroxyhexanoate and 5- (S)
hydroxyhexanenitrile
Candida boidinii E2: FDH (1.2.1.2) (S)-2-Amino-5-(1,3-dioxolan- Bristol-Myers Squibb,
or Pichia pastoris [E1: PheDH 2-yl)-pentanoic acid USA
From Thermoactinomyces
intermedius]
Trigonopsis D-Amino acid L-6-Hydroxynorleucine Bristol-Myers Squibb,
variabilis oxidase USA
ATCC 10679
Baker's yeast (= Reductase (R)-2,2,6- Hoffmann La-Roche,
Saccharomyces trimethylcyclohexane- CH
cerevisiae) 1,4-dione
Cryptococcus E1: Lactamase L-Lysine Toray Industries Inc.,

laurentii (3.5.2.11) and [E2: Japan
(E1) [and Racemase
Achromobacter
obae (E2)]a
Saccharomyces Pyruvate PAC ephedrine and Krebs Biochemicals
cerevisiae decarboxylase pseudoephedrine & Industries Ltd.,
India
Candida rugosa Enoyl-CoA hydratase R)--Hydroxy-n-butyric acid Kanegafuchi
and Chemical Industries
(R)--Hydroxy-isobutyric acid Co., Ltd., Japan
Rhodotorula rubra L-Phenylalanine L-phenylalanine Genex Corporation,
ammonia-lyase USA
Saccharomyces cerevisiae is the most thoroughly investigated eukaryotic microorganism and

Page 11
therefore the most frequently employed yeast strain in yeast-mediated whole-cell
biotransformations. Regarding classical approaches, S. cerevisiae was almost always among
the first biocatalysts chosen to solve the corresponding catalytic problem. However, finally
only three types of enzymatic reactions were successfully performed with S. cerevisiae whole
cells, namely oxidoreductase, hydrolase and lyase mediated biotransformations. The same is
true for alternative yeast strains which were found due to biodiversity screening approaches.
Alternative yeast strains employed for classical whole-cell biocatalysis are for example
Candida sp., Cryptococcus sp., Geotrichum sp., Issatchenkia sp., Kloeckera sp., Kluyveromyces
lactis, Pichia sp., Rhodotorula sp., Rhodosporidium sp., Schizosaccharomyces pombe,
Torulopsis sp., Trichosporon sp., Yarrowia lipolytica and Zygosaccharomyces rouxii. Yeast
strains employed for biotransformations in industry include Candida sp., Cryptococcus
laurentii, Geotrichum candidum, Pichia sp., Rhodotorula rubra, Saccharomyces cerevisiae,
Trigonopsis variabilis, and Zygosaccharomyces rouxii (Pscheidt & Glieder, 2008).
Biopharmaceuticals
Since the early 1980s, yeasts have been utilized for heterologous production of a variety of
proteins. The production of heterologous proteins in yeasts holds enormous potential for
biotechnological processes. A major breakthrough in heterologous protein expression in
yeast was the cloning, expression, processing, and secretion of human proinsulin in S.
cerevisiae in the 1980s (Branduardi and Porro, 2012). Yeasts are intensively being developed
as protein expression systems and in comparison to mammalian cell lines have higher
productivity, higher cell yields, shorter fermentation cycles, can be cultured in defined media
under relatively inexpensive conditions, can efficiently secrete proteins, possess
posttranslational modification pathways, nonpathogenic and non-pyrogenic (Demain &
Vaishnav, 2009).
Several yeast species have been investigated for heterologous protein production, namely
Saccharomyces cerevisiae, Pichia pastoris, Hansenula polymorpha, Kluyveromyces lactis,
Schizosaccharomyces pombe, Yarrowia lipolytica, Arxula adeninivorans (elik & alk, 2012,
Demain & Vaishnav, 2009; Porro et.al., 2011).
Recombinant proteins
Pharmaceuticals
Statins and other neutral products
Vitamins
Value of Product
Food Ingredients Antioxidants

Flavours
Antibiotics
Fine Chemicals Chiral building blocks
Enzymes
Ethanol
Fuels & Bulk Chemicals Solvents
Feed additives (amino acids)
Polymer building blocks
Volume of Product
Figure-3: Volume versus values of biotechnological products (Hong & Nielsen, 2012)

Page 12
The majority of biopharmaceuticals that have been approved for therapeutic applications by
regulatory authorities are proteins that have been produced by means of recombinant DNA
technology in various expression systems. They constitute approximately one-sixth of the
total pharmaceutical market and are its fastest growing segment. The biopharmaceuticals
are high value low volume products as shown in Figure-3 and their selling prices
recombinant proteins per gram are $375 for human insulin $23,000 for tPA $35,000 for
human growth hormone $384,000 for Granulocyte macrophage colony Stimulating factor
(GM-CSF) $450,000 for G-CSF $840,000 for Erythropoietin (EPO) compared to approx. $0.003
for bakers yeast (Demain & Vaishnav, 2009.). Table 6 shows examples of commercial
recombinant protein pharmaceuticals and vaccines produced by yeasts.
Table 6: Representative Commercial Recombinant Protein Pharmaceuticals and Vaccines

Produced in Yeasts (Johnson & Echavarri-Erasun, 2011)
Product Use Yeast Host
Short-acting recombinant insulin Diabetes mellitus Saccharomyces cerevisiae
Granulocyte macrophage colony Bone marrow transplantation S. cerevisiae
Stimulating factor (GM-CSF) Regulation of hematopoiesis S. cerevisiae
Acute myologenous leukemia
Hirudin/lepuridin Anticoagulant S. cerevisiae
Urate oxidase Hyperuricemia S. cerevisiae
Platelet-derived growth factor Diabetic ulcers S. cerevisiae
Human serum albumin Excipient, shock, cirrhosis, other S. cerevisiae
uses
Erythropoietin Renal disease S. cerevisiae
Glucagon Hypoglycemia S. cerevisiae
Human growth hormone Dwarfism, tissue repair S. cerevisiae
Platelet derived growth factor Anemia S. cerevisiae
Insulin Diabetes S. cerevisiae
Hepatitis A vaccine Hepatitis A S. cerevisiae
Hepatitis B vaccine Hepatitis B S. cerevisiae., Ogataea
polymorpha,
Komagataella pastoris
Diphtheria, tetanus, pertussis Hepatitis B S. cerevisiae
Haemophilus influenzae type B Combination vaccines and polio S. cerevisiae
Human papilloma virus (HPV) Human papillomavirus Kom. pastoris
antigen
In general, biopharmaceuticals are used to compensate for deficiency or lack of body

proteins important for normal functioning of the organism. They can be divided mainly into
the following categories: blood factors, thrombolytics and anticoagulants, hormones,
enzymes, growth factors, interferons and interleukins, vaccines and monoclonal antibodies.
Of the 211 biopharmaceuticals that have gained regulatory approval by the end of 2011, 66
(31 %) were produced in Escherichia coli, 31 (15 %) in yeast (of those, 30 in Saccharomyces
cervisiae and 1 in Pichia pastoris) and 91 (43 %) in mammalian cells. E. coli, yeast and
mammalian cells together account for the production of 89 % of approved
biopharmaceuticals and form the topic of this review (Martnez et al. 2012, Berlec &
Strukelj, 2013). Global market for recombinant protein drugs is 110 Billion USD at 2011 and
expected to rise to 160 Billion USD at 2014 (Martnez et. al., 2012).

Page 13
Metabolic Engineering in Saccharomyces cerevisiae
Metabolic engineering is the enabling science of development of efficient cell factories for
the production of fuels, chemicals, pharmaceuticals, and food ingredients through microbial
fermentations. The yeast Saccharomyces cerevisiae is a key cell factory already used for the
production of a wide range of industrial products. With the development of genetic
engineering in the 1970s, it became possible to produce compounds that are not native to
microbes, such as pharmaceutical proteins like human insulin and human growth hormone
using fermentation technology. Genetic engineering also allowed the transformation of
microbes into cell factories for the production of chemicals through so-called metabolic
engineering, a field dedicated to design of microbial metabolism to efficiently convert cheap
raw materials like glucose, sucrose, and biomass-derived sugars into fuels and chemicals
(Hong & Nielsen, 2012). Objective in Metabolic Engineering (Bettiga et.al., 2010): Extending
substrate range; metabolic engineering strategies aimed at endowing yeast with cellulolytic,
xylanolytic and amylolytic capacities, with a strong focus on S. cerevisiae and ethanol
production; Extending product range; introduction of novel pathways for the products
yeasts normally do not produce; Improving Cellular Properties: improving existing products,
redirecting carbon flux and reducing by product formation.
Volume
Saccharomyces cerevisiae
Glucose Biofuels
Sucrose Bulk chemicals
Galactose Fine chemicals
Xylose Protein drugs
Synthetic Biology Value
Systems Biology
Figure-4: Saccharomyces cerevisiae Cell Factory (Hong & Nielsen, 2012)
The rapidly expanding variety of chemicals produced by engineered S. cerevisiae, ranging

from commodity chemicals such as 1,2-propanediol, lactic acid, succinic acid through
ascorbic acid to fine chemicals such as resveratrol, or valancene (Table 7). The engineered
product range of S. cerevisiae encompasses compounds that are closely linked to primary
metabolism as well as molecules whose formation from yeast central metabolites requires
the functional expression of entire heterologous and/or synthetic pathways (Nielsen et.al.,
2013; Liu et.al., 2013). Isoprenoids, also referred to as terpenes or terpenoids, are among
the most diverse class of natural products, consisting of over 40 000 structurally different
compounds, which have been isolated from animal and microbial species as well as a wide
variety of plant organs. Isoprenoids form a good example of a class of compounds whose
production by metabolically engineered S. cerevisiae requires functional expression and/or
deregulation of multi-enzyme pathways leading from central metabolism to complex

Page 14
products (Wither and Keasling, 2007; Branduardi and Porro, 2012). Several lipid compounds
can be expressed in Saccharomyces cerevisia for the production of novel, high-value lipid
compounds. (Veen, M. and Lang, 2004)
Table 7: Example of products from S. cerevisiae (Hong & Nielsen, 2012, Buijs et.al., 2013)
Categories Products Categories Products
Biofuels Ethanol Fine -amyrin, -carotene
Biobutanol chemicals Amorpha-4,11-diene
Biodiesels Valencene and
Bisabolene (D2 dieselfuel, bisabolene) amorphadiene
The sesquiterpenoids:farnesene, amorphadiene Casbene (an anti-fungal
fatty acid ethyl esters (FAEEs) (biodiesel) diterpene)
Bulk 1,2-propanediol Cinnamoyl anthranilates
chemicals D-ribose and ribitol Cubebol, Linalool
L-lactic acid Eicosapentaenoic acid
Polyhydroxyalkanoates (EPA)
Pyruvic acid Farnese and geranyl
Succinic acid geraniol, L-ascorbic acid
Protein Insulin-like growth factor 1 (fhlGF-1) Methylmalonylcoenzyme

drugs Glucagon A
Single-chain antibodies (scFv) Patchoulol
Hepatitis surface antigen (HBsAg) Resveratrol
Parvovirus B19 VP2 Vanillin
Epidermal growth factor (EGF) Se-methylselenocysteine
Immunoglobulin G Non-ribosomalpeptides
Hepatitis B virus surface antigen (HBsAg)
L1 protein of human papillomavirus (HPV) type16
S. cerevisiae was among the first organisms to be designated generally recognized as safe
(GRAS), and the first genetically modified organism (GMO) used for recombinant production
of food and feed additives. In 1990, a genetically modified strain of S. cerevisiae became one
of the first GMOs to be approved for food-use in the UK. It is used in bakery products for
enhanced production of carbon dioxide. Many genes encoding desirable properties have
also been cloned in beer and wine yeast strains, S. pastorianus, and S. bayanus. However,
these strains have not been used commercially because of negative public perception
(Dequin, 2001; Attfield et.al, 2003; Randez-Gill et.al., 2003; Donalies et.al., 2008).
Oleaginous Yeasts or Single Cell Oils

The ability of certain microorganisms to accumulate high amounts of lipids has been known
for years (see Figure-5). Oily yeasts have been described to be able to accumulate lipids up
to 20% of their cellular dry weight. These yeasts represent a minor proportion of the total
yeast population, and only 5% of them have been reported as able to accumulate more than
25% of lipids. The oily yeast genera include Yarrowia, Candida, Rhodotorula,
Rhodosporidium, Cryptococcus, Trichosporon, and Lipomyces. More specifically, examples of
oleaginous yeasts include the species: Lipomyces starkeyi, Rhodosporidium toruloides,
Rhodotorula glutinis, and Yarrowia lipolytica. Yeast do exhibit advantages for lipid
production over other microbial sources, namely, their duplication times are usually lower
than 1 h, are much less affected than plants by season or climate conditions, and their
cultures are more easily scaled up than those of microalgae. Additionally, some oily yeasts
Page 15
have been reported to accumulate oil up to 80% of their dry weight and can indeed generate
different lipids from different carbon sources or from lipids present in the culture media.
Thus, they can vary their lipid composition by replacing the fatty acids present in their
triglycerides. Due to the diversity of microorganisms and growth conditions, oily yeasts can
be useful for the production of triglycerides, surfactants, or polyunsaturated fatty acids
(Ageitos et.al, 2011; Papanikolaou and Aggelis, 2011a,b; Christophe et.al., 2012).
Figure-5: Lipid accumulation in yeast (Ageitos et al., 2011)
Table 8: Lipid Accumulation of Selected Oleaginous Yeasts

(Ageitos et al., 2011)
Species Lipid accumulation
(% D.W.)
Cryptococcus curvatus 69
Cryptococcus albidus 65
Candida sp 107 42
Lipomyces starkeyi 73
Rhodotorula glutinis 72
Rhodosporidium toruloides 76
Rhodotorula graminis 36
Trichosporon pullulans 65
Yarrowia lipolytica 36
Of the 600 species of yeast, only 30 have been characterized as being able to accumulate
more than 25% of their dry weight as lipids. More specifically, examples of oleaginous yeast
species include: Cryptococcus albidus, Lipomyces lipofera, Lipomyces starkeyi,
Rhodosporidium toruloides, Rhodotorula glutinis, Trichosporon pullulan, and Yarrowia
lipolytica, (formerly classified as Candida lipolytica) (Li and Liu, 2008; Ageitos et.al., 2011)

Page 16
(Table 8). Additionally, there are some other oleaginous yeast species feasible for oil
production and thus possible biodiesel producers (Leiva-Candia et.al., 2014).
Psychrophilic or Cold Adapted Yeasts

More than 80 % of the Earths environments exhibit temperatures below 5 0C. The major
fraction of cold habitats is represented by deep oceans, Arctic and Antarctic snow- and ice
caps, permafrost soils, sea ice and high mountain glaciers. Antarctica is the most
representative example of a terrestrial cold habitat, of which almost the totality of surface is
covered by ice and snow. In many ecosystems, cold conditions are frequently associated
with other limiting factors (e.g. low water availability (Aw) and nutrient availability, high
hydrostatic pressure and oxidative stress, high solar irradiation) which make such extreme
habitats very inhospitable (or even life-limiting) ecosystems (Buzzini et.al., 2012)
Yeasts may be better adapted to low temperatures than bacteria. Yeasts belonging to genera
such as Bullera, Candida, Cryptococcus, Cystofilobasidium, Debaryomyces, Kondoa,
Leucosporidium, Metschnikowia, Mrakia, Pseudozyma , Rhodotorula, Sakaguchia,
Sporopachydermia, Sympodiomyces and Trichosporon have been identified in various
habitats of Antarctica. Psychrophilic yeasts have an optimum temperature for growth at
about 15 C or lower, a maximum up to 25 C but are still capable of growing at 0 C or
below. Low temperature adaptation is associated with the proportion of unsaturated fatty
acids in composition of the membranes. The biotechnology of cold-adapted yeasts is not
merely confined to the production of cold-active enzymes: a number of recent studies
carried out at the laboratory scale have demonstrated that their future perspectives could
also be directed towards the production of polymeric compounds. The heterologous
expression of proteins from cold-adapted yeasts in suitable hosts is an additional possible
venture. The use of cold-adapted yeasts for agricultural, food and environmental
biotechnologies could represent another attracting way to ensure their possible economical
exploitation: low-temperature wine-making and beer-making, control of post-harvest
diseases of fruits and seeds, and bioremediation of hydrocarbon polluted cold ecosystems
represent undoubtedly the most relevant challenges. Cold-adapted spoilage yeasts can
potentially cause physical and chemical deterioration of foods, thus determining a potential
risk to consumers and a relevant economic burden for food (Buzzini et.al., 2012).
Table 9: Examples of Psychrophilic Enzymes and Proteins from Cold Adapted Yeasts (Buzzini
& Margesin, 2014. Margesin & Feller, 2010).
Products Yeast Genera or sp. Applications
Antifreeze proteins (AFPs) Glaciozyma antarctica, food industry (ice cream, frozen
Glaciozyma sp., and dough, fruits, and vegetables to keep
Rhodotorula glacialis longer shelf life cryopreservation of
Rec-Saccharomyces cerevisiae red blood cells, cryosurgery
Amylases, Xylanases, Food industry
Chitinases, and Lysozymes
Lipases Candida antarctica Food industry, pharmaceuticals or
cosmetics
Pectinolytic Enzymes Cystofilobasidium, fruit-processing ndustry, reducing
Cryptococcus, juice viscosity and for improving the

Page 17
and Mrakia clarity
-galactosidases Guehomyces lactose-free milk and sweet whey

Proteases Glaciozyma antarctica additives in detergents and foodstuffs
bioremediators, in iotransformations,
and in molecular biology maturation of
slow-ripening cheeses in a low-
temperature and low-moisture
environment
Phytases and Esterases Cryptococcus laurentii Animal feed
Rhodotorula mucilaginosa biotransformations
Yarrowia lipolytica
Cold-adapted organisms, thriving permanently at near-zero temperatures, synthesize cold-

active enzymes to sustain their cell cycle as shown in Table 9. These enzymes are already
used in many biotechnological applications requiring high activity at mild temperatures or
fast heat-inactivation rate. The useful applications of these enzymes are widespread to a
large number of industries like textile industry, food and dairy industry, brewing and wine
industry, laundry, etc. Cold-active hydrolytic enzymes like lipases, proteases, cellulases, and
amylases can be used as an active agent in detergents applied for cold washing. Other
potential applications of psychrophilic enzymes, apart from these, are in processes such as
the hydrolysis of lactose in milk using -galactosidases, extraction and clearing fruit juices
using pectinases, meat tenderization or taste improvement of refrigerated meat using
proteases, betterment of bakery products using glycosidases (e.g., amylases, xylanases).
lipases are some of the most useful enzymes, especially in the production of high added
value substances, such as fine chemicals, pharmaceuticals, foods, polymers, detergents,
biodiesel. Among all known lipases from different environments and organisms, of special
importance are cold-active lipases A and B from the yeast Pseudozyma (Candida) antarctica,
which have been thoroughly researched.
Anti-freeze proteins (AFPs) are a useful material in commercial applications such as food
industry, cryosurgery, and cryopreservation of red blood cells, oocytes, organs, and tissues.
Food industry is a significant field where antifreeze proteins can be applied. AFPs can be
added to various food items, such as ice cream to make a better texture and to frozen
dough, fruits, and vegetables to keep longer shelf life (Buzzini & Margesin, 2014).
At the industrial level, the best-known representative of polar microorganisms is certainly

the yeast Candida antarctica , as its species name unambiguously refers to the sampling
origin. This yeast produces two lipases, A and B, the latter being sold for instance as
Novozym 435 by Novozymes (Denmark). Although the moderate heat-stability of this lipase
in aqueous solutions can be of concern, this enzyme is stabilized in its immobilized form. As
a result of its substrate and stereospecificity, lipase B is involved in a very large number of
organosynthesis applications related to food/feed processing, pharmaceuticals or cosmetics.
In a survey of patents related to Antarctica it was shown that lipases from C. Antarctica by
far dominate the number of process- or product-based patents. This is a significant example
of the potential for novel catalysts from genetic resources in cold environments. The market
for enzymes used in detergents (Margesin and Feller, 2010)
Agricultural and Biocontrol Applications

Page 18
Postharvest losses of stored fruits, vegetables and grains due to decay by fungal pathogens
can be very significant, and in fact, addressing this problem is receiving increasing
importance as worldwide demand for food increases. Postharvest decay of fruits can be
reduced by avoiding injury to the fruit during harvest and subsequent handling, stringent
sanitation practices, the use of chemical fungicides during cold and/or modified atmosphere
storage. These beneficial practices, however, are usually not enough to sufficiently protect
harvested fruits from spoilage, caused mainly by several fungal species belonging to
Alternaria, Aspergillus, Botrytis, Fusarium, Geotrichum, Gloeosporium, Mucor, Monilinia,
Penicillium, Rhizopus and other genera. There is an increasing concern about the
environmental effects and safety of chemical pesticides and fungicides all over the world.
Traditionally, the postharvest diseases are controlled by fungicides. The main problems
involved in fungicide utilization are related to environmental pollution and public health
concerns (Pimenta et. al., 2009).
Among the alternative approaches that have been explored, the use of microbial antagonists
like yeasts, fungi and bacteria has been demonstrated to be quite promising and gained
increased attention. The majority of microbial organisms studied for their potential use as
postharvest biocontrol agents belongs to yeasts due to the fact that many of these
microorganisms are considered as generally recognized as safe sa shown in Table 10.
Features required for any effective BCA should include a rapid growth rate in fruit wounds,
the effective utilization of the nutrients present in the wound, and the capability to survive
and develop at the infection site better than the pathogen. And to do so all, this should
occur at a low temperature, acidic pH, conditions of osmotic stress, resistance to
desiccation, tolerance to chemicals, The yeast in postharvest biocontrol formulations
apparently presents advantages over other organisms: easy to cultivate, fast growing and
readily found in a variety of substrates and conditions.
Table 10: Examples of Yeast Antagonists Applied Preharvest to Reduce Postharvest

Pathogens (Schisler et.al, 2011)
Yeast/Yeast-like Fungus Disease Assessment Utilized
Aureobasidium pullulans Various postharvest rots Cherry fruit
Aureobasidium pullulans and Gray mold, blue mold, bulls eye rot Apple fruit
Rhodotorula glutinis Blue mold Apple fruit
Candida sake Gray mold Strawberry fruit
Cryptococcus albidus Blue and gray mold, side rot Pear fruit
Cystofilobasidium infirmominatum, Various postharvest rots Cherry fruit
Cryptococcus laurentii, and Gray mold, other postharvest rots Table grape fruit
Rhodotorula glutinis Anthracnose Mango fruit
Cryptococcus laurentii
Metschnikowia fructicola
Rhodotorula minuta
Research on the use of yeasts as BCAs has mainly focused on their use for managing
postharvest diseases, mainly of fruit; however, this application represents only a small
portion of the complete spectrum of plant disease management . Only six products based on
different yeast species have been registered for postharvest use (Buzzini and Margesin,
2014):
Page 19
1. Aspire, based on C. oleophila (Ecogen Inc., Langhorne, USA);
2. Yield Plus, based on C. albidus (Lallemand, Montreal, Canada);
3. ShemerTM, based on Metschnikowia fructicola (Bayer CropScience);
4. Candifruit, based on Candida sake (Sipcam-Inaagri, SA Valencia, Spain);
5. Nexy, based on C. oleophila (Lesaffre-Bionext, France); and
6. Boni-Protect, based on A. pullulans (Biofa, Mnsingen, Germany)
The scarcity of commercial products for postharvest use has been discussed and the main
problems mentioned are the small number of companies involved in the development of
biological products, the small size of the postharvest market, the expense and length of time
required for selection and registration (Buzzini and Margesin, 2014).
Environmental Applications
Yeasts have important roles in agriculture as agents of biocontrol, bioremediation, and as
indicators of environmental quality. Especially basidiomycetous yeasts have important roles
in the biotransformation and degradation of pollutants and xenobiotics. This activity is
thought mostly to be due to the production of solubilizing acids within microbial consortia
such as biofilms. Non-pathogenic species of Cryptococcus and Trichosporon are capable of
degrading polysaccharides, phenolic compounds, complex organic acids, and C2
hydrocarbons. Strains of Pseudozyma have been reported to degrade plastics and related
compounds (Pimenta et.al., 2009; Middelhoven, 2009; Seo et.al., 2007). The coloured
recalcitrant compounds can be removed from molasses based wastewaters by yeast
Issatchenkia orientalis which is main problem in bakers yeast industry (Tondee et.al., 2008).
Basidiomycetous yeasts were shown to degrade starch, pullulan, dextran, xylan,

polygalacturonate, galactomannan, and tannic acid as sole carbon source. They were unable
to grow on cellulose, chitin, arabinogalactan, and xanthan gum. Several basidiomycetous
yeasts grow on phenolic compounds. Basidiomycetous yeast diversity varied in 54 different
soils varieties under different management regimes, including species of Cryptococcus,
Rhodotorula, and Trichosporon. These results indicate that basidiomycetous yeasts are
involved in the natural biodegradation of biomass and may have potential for remediation of
various industrial waste streams. Table 11 shows representative examples of treatment of
recalcitrant chemicals and pollutants by different yeast species.
Table 11: Examples of Treatment/Biotransformation of Recalcitrant Chemicals, Pollutants,

and Xenobiotics by Yeast (Middelhoven, 2009; Gonalves et.al, 2009; Begum et.al., 2003;
Tondee et.al., 2008; Yang et.al, 2013; Krallish et.al, 2006; Seo et.al., 2007; Bleve et.al.,
2011; Watanabe et.al., 2009; Rossi et.al, 2009)
Waste/Compounds Yeast
Phthalic Acid Esters Saccharomyces cerevisiae
phenol, acetophenone, acetone, amethylstyrene, Trichosporon cutaneum R57
benzoic acid, dimethyl phenyl carbinol, methanol and
isopropylbenzene
Olive mill wastewater Candida rugosa, Candida cylindracea and Yarrowia
lipolytica
Decolorization of molasses wastewater Issatchenkia orientalis
Phenol Rhodotorula creatinivora
Cryptococcus terreus (Cold-adapted)
Plastic waste Pseudozyma jejuensis
Page 20
Phosphorus removal Hansenula anomala
Nitrate assimilation Hansenula polymorpha
Removal of lactose/other sugars from wastewater Kluyveromyces marxianus
Due to the ability of various yeast species to grow on a wide diversity of substrates, including
aromatic molecules, alkanes, lipid compounds, amines and other recalcitrant compounds,
they have the capacity for the transformation of deleterious compounds to innocuous
derivatives. Y. lipolytica transformed 2, 4, 6-trinitrotoluene (TNT) to derivative products.
Molecular oxygen and oxidative enzymes such as cytochrome P450s and oxygenases are
often involved in the degradation pathways. Phenolic degradation is also carried out by
several yeasts, particularly basidiomycetous genera such as Cryptococcus, Rhodotorula, and
Trichosporon. Several species of Candida and Trichosporon spp. are recognized to efficiently
transform halogenated aromatic compounds. Polycyclic aromatic hydrocarbons were
degraded by species of Candida, Cryptococcus, Rhodotorula, and Trichosporon. Recombinant
yeasts expressing a soybean cytochrome P450 enhanced the metabolism of phenylurea
herbicides. It has been reported that Pseud. jejuensis is capable of degrading certain plastic
wastes (Seo et.al., 2007). Immobilized and cold-adapted yeast systems are being developed
as industrial biodegradative systems. Yeast systems have also been developed for the
sensing and accumulation of various ecotoxicants such as heavy metals. Xanthophyllomyces
dendrorhous and Ph. rhodozyma have been reported to degrade ochratoxin, a potent
mycotoxin. Trichosporon spp. are capable of degrading ochratoxin A and zearalenone in
laboratory cultures. S. cerevisiae strains have been reported to degrade patulin during cider
fermenations. The ability of yeasts to degrade mycotoxins has important health implications,
since these toxicants occur in commodities, such as feeds, foods, and beverages, and are
considered as serious health hazards (Schisler et.al, 2011).
Table 12: Applications of Y. lipolytica in the treatment/upgradation of wastes and

use of wastes as inexpensive alternative substrates (Bankar et.al., 2009)
Waste Effect of treatment Other products/applications
Olive mill wastewater 80% COD reduction Single-cell protein, lipases
Olive mill wastewater 1.47% to 41% COD reduction Lipases, citric acid
Olive mill wastewater 80% COD reduction Lipases
Olive mill wastewater 22% to 52% COD reduction Lipases
Oil waste water 82% COD reduction
Palm oil mill effluent 95% COD reduction
Fish wastes solid state 29% fat reduction Fish meal
Yarrowia lipolytica is a ascomycetous yeast species with distinctive physiological features

and biochemical characteristics that are significant in environment-related matters. Strains
naturally present in soils, sea water, sediments and waste waters have inherent abilities to
degrade hydrocarbons such as alkanes (short and medium chain) and aromatic compounds
(biphenyl and dibenzofuran). The natural ability of the yeast to produce surfactants can also
help in remedial procedures. Apart from the aliphatic and aromatic hydrocarbons that it
effectively breaks down, wild-type and recombinant strains can also degrade nitroaromatic,
halogenated and organophosphate compounds. The yeast has an inherent ability to detoxify
metal ions and this has been exploited for developing bioremedial applications involving
biosorption and for the synthesis of nanoparticles. The organism can act on different kinds of
Page 21
wastes including oily and solid wastes and bring about a reduction in COD or yield value-
added products such as citric acid, surfactants, enzymes and single cell oil. Some applications
of Y. lipolytica are shown in Table 12 in the treatment/upgradation of wastes and use of
wastes as inexpensive alternative substrates. Moreover, some strains have been
exceptionally versatile with respect to their interactions with pollutants and may be
projected as major players in the development of technologies in the future (Bankar et.al.,
2009; Nicaud, 2012; Zinjarde et.al., 2014).
Conclusions
Yeasts are remarkable organisms and they are more than Saccharomyces cerevisia used in
traditional fermentations. Their involvement and importance in traditional food
fermentations are unparalled by other organisms of biotechnological relevance. The number
of yeast species identified and their biotechnological potential are accelerating due to a
number of properties and developments. New developments in engineering pathways in
yeast using sytem biology and rDNA technologies have introduced novel capabilities to
extend substrate range, to improve cellular properties and to produce new products yeast
can not produce. S. cerevisiae and certain other yeast species are being developed for the
production of biofuels from cellulosic materials and potentially other substrates. During the
past decade, yeast systems have been developed for the production of heterologous
proteins in high yields. Yeasts are increasingly important as sources of biocatalysts for the
production of comodity and high-value fine chemicals and enzymes. Many species, especially
basidiomycetous yeasts have strong oxidative metabolism that enables the bioremediation
and degradation of chemicals, pollutants such as aromatic compounds, plastics, and other
recalcitrant compounds and polymers. These advances provide important strategies and
tools for enhancing the biotechnological importance of yeasts.
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Yeast Biotechnology: Diversity and

Conference Paper April 2014

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Role of Yeast in Traditional Food Fermentations

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Figure-1: Traditional Uses of Yeasts (adapted from Walker, 1998)

Food and Feed Yeasts

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Modern Applications of Yeasts

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Table 1: Biotechnologically Important Yeast sp. (Johnson, 2013a,b)

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Figure-2: Modern Applications of Yeasts (adapted from Walker, 1998)

Food and Feed Enzymes from Yeasts

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Table 2: Industrial Enzymes from Yeasts (Johnson & Echavarri-Erasun, 2011)

Table 3: Examples of Industrial Recombinant Enzymes

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Yeast Metabolites and Bulk and Fine Chemicals

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Biocatalysis and Fine Chemicals

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Table 5: Industrial biotransformations employing wild type yeast whole-cell biocatalysts

Cryptococcus E1: Lactamase L-Lysine Toray Industries Inc.,

Saccharomyces cerevisiae is the most thoroughly investigated eukaryotic microorganism and

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Food Ingredients Antioxidants

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Table 6: Representative Commercial Recombinant Protein Pharmaceuticals and Vaccines

In general, biopharmaceuticals are used to compensate for deficiency or lack of body

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Sucrose Bulk chemicals

Galactose Fine chemicals

Xylose Protein drugs

Synthetic Biology Value

The rapidly expanding variety of chemicals produced by engineered S. cerevisiae, ranging

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Protein Insulin-like growth factor 1 (fhlGF-1) Methylmalonylcoenzyme

Oleaginous Yeasts or Single Cell Oils

Figure-5: Lipid accumulation in yeast (Ageitos et al., 2011)

Table 8: Lipid Accumulation of Selected Oleaginous Yeasts

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Psychrophilic or Cold Adapted Yeasts

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

-galactosidases Guehomyces lactose-free milk and sweet whey

Cold-adapted organisms, thriving permanently at near-zero temperatures, synthesize cold-

At the industrial level, the best-known representative of polar microorganisms is certainly

Agricultural and Biocontrol Applications

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

Table 10: Examples of Yeast Antagonists Applied Preharvest to Reduce Postharvest

Basidiomycetous yeasts were shown to degrade starch, pullulan, dextran, xylan,

Table 11: Examples of Treatment/Biotransformation of Recalcitrant Chemicals, Pollutants,

Table 12: Applications of Y. lipolytica in the treatment/upgradation of wastes and

Yarrowia lipolytica is a ascomycetous yeast species with distinctive physiological features

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

27th VH Yeast Conference, April 1 4th 1 5th, 2014, Istanbul.

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