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ALBERT K. ATHNASIOS, Fleischmanns Yeast, Division of Burns Philp Food Company,

Fenton, United States
MICHAEL QUANTZ, Research Institute for Bakers Yeast, Berlin, Germany

1. Introduction . . . . . . . . . . . . . . . . . . . . 1 3.2.6. Wine Yeasts . . . . . . . . . . . . . . . . . . . 11

2. The Yeast Cell . . . . . . . . . . . . . . . . . . . 2 3.2.7. Distillers Yeasts . . . . . . . . . . . . . . . . 11
3. Food and Feed Yeast . . . . . . . . . . . . . 3 4. Yeast-Derived Products . . . . . . . . . . . 11
3.1. Chemical composition of Yeast . . . . 3 4.1. Flavor Products . . . . . . . . . . . . . . . . 11
3.2. Production of Biomass . . . . . . . . . . 4 4.2. Nutritional Yeast . . . . . . . . . . . . . . . 12
3.2.1. Bakers Yeast . . . . . . . . . . . . . . . . . . 4 4.3. Colorants Derived from Yeast . . . . . 12
3.2.2. Active Dry Yeast (ADY) and Instant 4.4. Yeast-Derived Enzymes . . . . . . . . . 13
Active Dry Yeast (IADY) . . . . . . . . . 9 4.5. Cosmetic and Pharmaceutical
3.2.3. Candida utilis (Torula) Yeast . . . . . . . 10 Products . . . . . . . . . . . . . . . . . . . . . 13
3.2.4. Dairy Yeasts . . . . . . . . . . . . . . . . . . . 10 4.6. Yeast Engineering . . . . . . . . . . . . . . 14
3.2.5. Brewers Yeast . . . . . . . . . . . . . . . . . 10

1. Introduction [1] goods. Preparation of compressed brewers

yeast began in the late 1700s in Great Britain
The first records on the use of yeast date back to and Austria, where the aeration of yeast to
6000 B.C. Egypt, where a type of acid beer called increase the yeast yield was invented in 1835
boozah was produced. by MAUTNER.
The production of wine, beer, and leavened In 1868 FLEISCHMANN founded the com-
bread probably developed over the next few pressed bakers yeast industry in the United
thousand years. In Roman times, beer was a States. Concurrent improvements in under-
common drink in Spain and Germany. The standing the biochemistry of yeast assimilation
German beer regulation (Bier-Reinheitsge- and fermentation ability and technical process
bot), coming into force in 1516, described a engineering greatly aided the development of
fermenting agent on barley malt in water, al- the modern yeast factory, dedicated for bakers
though yeast was not identified as a living yeast production.
organism at that time. Modern technology for biomass production
The nature of yeast long remained unknown (! Biotechnology, 1. General) dates from
until VAN LEEUWENHOEK developed the first World War I. Large-scale fermenters with
microscope and in 1860 PASTEUR established effective aeration devices were developed and
the role of yeast in the fermentation of sugar new continuous processes for effective product
to alcohol beyond doubt. HANSEN established separation by centrifugation were used. New
methods to isolate and breed pure cultures of raw materials (molasses, sulfite liquor contain-
yeast strains at the end of the 19th century. ing wood sugar, whey, starchy wastes from
The technology of microbial cell production potato processing, date syrup, etc.) were used.
on a commercial scale for food and feed has The number of yeast strains and species grown
developed only within the past 150 years. In on different raw materials also increased.
ancient times people recovered top-fermenting Four species of yeasts are presently used in
yeasts of the genus Saccharomyces from the foods: Saccharomyces cerevisiae, Saccharomy-
production of fermented beverages such as beer, ces uvarum (formerly Saccharomyces carlsber-
and used them as leavening agents in baked gensis), Candida utilis (Torula utilis), and

 2012 Wiley-VCH Verlag GmbH & Co. KGaA, Weinheim

2 Yeasts

Kluyveromyces fragilis (formerly Saccharomy- properties are often very similar, and separate
ces fragilis). names have not been justified taxonomically.
S. cerevisiae and S. uvarum, the yeasts of the For various industrial processes, special
brewing industry, are byproducts of beer pro- characteristics of production strains are of great
duction, so-called secondary yeasts. Other importance; therefore, strain differentiation
yeasts are grown especially for food and feed may be more valuable than species
protein use and are often called primary yeasts: classification.
T. utilis is grown in sulfite liquor, and K. fragilis
on cheese whey.
Detailed chemical analysis shows that yeast 2. The Yeast Cell
is a rich source of protein, amino acids, essential
minerals like chromium, selenium, and zinc, It is difficult to define yeast precisely on the
and one of the richest sources of B group basis of general morphological and physiologi-
vitamins. cal considerations. For example, most yeast
Although fermentation and leavening of cells are colorless and transparent, while
doughs are the most common uses of yeast in some produce carotenoid pigments (e.g.,
food production, the application of yeast and Rhodotorula). Bud formation is widespread, but
yeast derivatives for nutritional fortification (as is not common to all yeasts. Cell shape, in most
protein and vitamin sources) of processed foods cases, is round to oval, with a diameter of
and also for refining their flavor profile is 47 mm. Several species of some genera may
growing. produce a true mycellium.
After the Saccharomyces yeasts genome was Electron micrographs of thin sections of
sequenced in 1996, the understanding of the yeast cells show the existence of a cell wall, a
functionality of yeast genes has reached a state plasma membrane, nucleus, mitochondria, and
where yeast and its metabolism can be engi- vacuoles.
neered at high rates and yields. Figure 1 shows a bakers yeast cell cross
The taxonomy of about 750 yeast species is section scheme with cell wall and internal
in a continual state of flux. As new microorgan- structures.
isms are discovered, increasing knowledge The cell wall makes up 15 to 20% of
about them results in the yeasts being shifted the dry weight of the cell. The proportional
within subgroups. However, such changes are weight of the cell wall usually decreases during
slow to be put into practice, except by the growth phase and increases again in the
taxonomists. stationary phase. The main cell wall compo-
Frequently, species were named on the basis nents are the polysaccharides glucan and man-
of the fermentation they were associated with, nan, chitin, protein, and lipids. Acid hydrolysis
e.g., Saccharomyces vinii or Saccharomyces of the cell wall results in the formation of
sake. The morphological and physiological glucose, mannose, glucosamine, amino acids,

Figure 1. Cross-section scheme of Bakers yeast (courtesy of J. HEINISCH, University of Osnabruck)

Yeasts 3

phosphates, and lipids. The chemical composi- and occasionally they contain degrading intra-
tion of the cell wall of S. cerevisiae shows great cellular substances.
variability, with carbohydrates varying from 60
to 91%, proteins from 6 to 13%, and lipids from
2 to 5.8%. The two main polysaccharides, glu- 3. Food and Feed Yeast [2]
can and mannan, are present in about equal
amounts. Glucans form the matrix of high- The world yeast market volume of about
molecular-mass polymers, from b- (13 and 4.5109 t in 2010 comprises 60% feed yeasts
16)-linked glucose monomers, embedding from several sources, 25% bakers yeast, and
chitin fibers, a polymer of b-(14)-linked about 5% brewers yeast. The remaining 10%
N-acetylglucosamine, which provide the enor- include yeast-derived products like yeast auto-
mous flexibility of the yeast cell wall. lysates, extracts, nutritional, and special yeasts
The cell-wall protein contains all the com- for wine and distillery.
mon amino acids, with high proportions of The most important yeast-fermented food
glutamic and aspartic acids. In contrast to the products are baked goods, beer, and wine, which
whole-cell protein amino acid composition, constitute a significant percentage of the human
cell-wall protein has a high content of sulfur diet. Inactive dried yeasts are sold for human
amino acids (cysteine and methionine) and, consumption because of their nutritional values:
especially in younger cells, contains SH Yeasts contain B group vitamins, folic acid,
groups. In older cells the protein chains are pantothenic acid, and selenium and chromium
linked by disulfide bonds (cystine). The struc- as essential trace elements. Additionally, the
tural protein of the cell wall is mainly bound to b-(13 and 16)-linked glucans from the yeast
polysaccharides to form a complex manno- cell walls have been considered as immunopro-
protein structure in which glucosamine acts as tective factors in recent times. The dried pow-
the connecting link between the polysacchar- ders or flakes are usually consumed by blending
ides and proteins. them with fruit juices or by sprinkling over
The lipid content of the cell wall varies both cereals.
with the species and growth conditions. Pub- The main feed, protein-rich yeast volumes
lished data show large variations, probably due are primarily grown on pulp and paper mill
to incomplete removal of the lipid-rich plasma waste liquors containing fermentable sugars.
membrane. Some of the lipid content is firmly Most of the brewers excess yeast is used in
bound to the cell wall and may play a role in feed as well, usually by co-drying it with spent
maintaining its orderly structure. C16:1, C18:1, grains, another byproduct of the brewing indus-
and C16:0 acids are the dominant fatty acids of try. But a considerable percentage is used in the
the cell-wall lipids. The plasma membrane production of yeast extracts or sold as debittered
lipids have a higher content of C18:1 acids; in brewers yeast for nutritional purposes.
addition they are enriched in phosphatidylino- In the production of distilled beverages, the
sitol and phosphatidylserine components of yeast is co-dried with the distillers solubles or
phospholipids compared to the cell lipids. the spent grain (DDGS) and used as feed. Active
The nucleus of yeast is surrounded by an dry yeast is also used as a supplement to rumi-
envelope characterized by the presence of many nant feeds and has been reported to increase feed
pores. efficiency in cattle.
Mitochondria are abundant in all fungi. Their
most morphological characteristics are the pres-
ence of inner and outer membranes. Size, shape, 3.1. Chemical composition of Yeast
number, and composition of mitochondria vary
widely under different growth conditions. The gross composition of yeasts is quite vari-
Vacuoles are important subcellular able. The percentage of proteins and nucleic
organelles in yeasts. Both vegetative and acids depend on the substrate and growth rate.
reproductive cells contain vacuoles that vary The yeast composition is outlined here on the
significantly in size. The vacuoles contain a basis of S. cerevisiae. The other two nutritional
variety of hydrolytic enzymes such as proteases, yeasts, namely C.utilis and Kluyveromyces
4 Yeasts

marxianus, have a somewhat similar concentrations of potassium and phosphorus

composition. in dry yeast are higher than those of other
elements such as Ca, Mg, and S, the mineral
Protein. The protein of yeasts is usually contributions made by an allowable daily
determined by Kjeldahl nitrogen determination serving of 20 g of dry yeast are minor consid-
and multiplication of the nitrogen content by ering the high mineral content in the bulk of an
6.25 (N6.25). The total nitrogen includes the average human diet.
nitrogen of the nucleic acids and, therefore, this However, the accumulation of micronutri-
procedure overestimates the protein content. ents like chromium and selenium in yeast are of
Protein calculated in this manner should be special interest. Specific active metal ion trans-
designated crude protein. A factor of N5.5 is porters were identified in the yeast plasma
more appropriate to estimate pure protein. membrane for iron and copper. The Cr uptake
Bakers yeast contains 69% nitrogen of mechanism is still under examination.
yeast dry solids (YDS). High nitrogen levels in There are several factors that restrict Se
the yeast result in higher dough raising power uptake by yeast under conventional batch prop-
but lower stability, both for compressed and agation conditions primarily because of the
active dry yeasts. toxic effects of Se salts on yeast growth. How-
Total yeast proteins are generally higher in ever, a new procedure for the propagation of
lysine and deficient in sulfur amino acids, me- food-grade Se-rich yeast has been developed,
thionine, cysteine, and cystine as compared to based on the concept that, under conditions of
cell-wall protein amino acid composition (see sulfur deficiency, selenium could replace sulfur
Chap. 2). The amino acid composition of yeast in yeast. The growth medium is fed incremen-
resembles that of oil-seed proteins, particularly tally so that the Se concentration never reaches
soy protein. toxic levels.

Vitamins. Yeast predominantly contains the Lipids. The lipid content of yeasts can vary
vitamins of the B complex (B1, B2, and B6), and between 4 and 7% (dry basis). About 1% can be
is thus an excellent source of B vitamins for extracted directly by solvents. The Weibull
human and animal nutrition. Stoldt determination of total lipids requires acid
However, yeast does not provide vitamin C hydrolysis prior to solvent extraction. Major
and fat-soluble vitamins such as A, E, K, and D. lipid constituents are fatty acid glycerides with
Nevertheless, dry yeast can function as a valu- a predominance of palmitic and oleic acids,
able vitamin source when used in combination sterols, and lipolipids.
with other food ingredients, as is generally the
case with human diets. Carbohydrates. Total carbohydrates
Bakers yeast contains 710% ergosterol, account for about 3035% of the yeast cell
mainly in the membranes. It serves as a precur- (dry basis). They consist mainly of carbohy-
sor for vitamin D2. Conversion occurs when drate storage compounds such as glycogen
ergosterol is irradiated with ultraviolet light. (a-glucan), the disaccharide trehalose, and the
This process competes with that of synthetic structural materials of the cell wall: the
vitamin D2, which is less expensive, but yeast b-glucans and mannans. Level of fiber in
treated in this way can still raise the dough, whole bakers yeast cells is about 18% (dry
enabling bread producers to have an in-situ weight basis).
fortification with vitamin D. Bakers yeast does
not contain 7-dehydrocholesterol, the precursor
of vitamin D3. 3.2. Production of Biomass [3]

Minerals and Trace Elements. Yeasts take 3.2.1. Bakers Yeast

up substantial quantities of macro- and micro-
nutrients from the surrounding growth medi- The production of bakers yeast, exclusively for
um. The mineral-ash content amounts to ca. the production of bread doughs, dates from the
8% on a dry solid basis. Although the late 1800s. It started with the production on
Yeasts 5

grain mashes, which were later replaced with 3. Active dry yeast: Progress was made in the
the least expensive source of fermentable sugar, late 1930s in the drying of compressed yeast
molasses. (CY) to the more stable active dry yeast
During the 20th century the major advances (ADY). Continued progress in the drying
in the production of bakers yeast have been the methods has led to ADY and instant active
following: dry yeast (IADY, 1970), replacing CY in
several bakery applications, particularly in
1. Fed-batch process (Zulaufverfahren): The industrial baking and in pizzerias.
use of incremental feeding was introduced to 4. Automation: During the past decades there
bakers yeast production by Danish and has been a rapid shift to automatic control of
German scientists between 1910 and 1920. the fermentation process. Parameters to be
This process is still used today, because it is controlled are the media flow, the pH, the
the only practical method that permits soluble oxygen, and the ethanol content of
production of yeast biomass without simul- the wort. The wort foam level is needed to
taneous production of large quantities of control risks of spill and is coupled to
ethanol (to avoid the Crabtree effect). antifoam-agent dosage.
2. Molasses: Before 1920 the mash consisted of
corn, malt, and malt sprouts. During the Figure 2 shows the block flow sheet of the
1920s and 1930s, the grains were slowly production process of bakers yeast with the
replaced with molasses as carbon and energy main educts and the products, cream yeast, fresh
source for yeast growth. yeast, and IADY.

Figure 2. Block flow sheet of the production of bakers yeast

6 Yeasts

Cane or beet molasses supplies sugars as a with roots blowers or dynamic, partially self
carbon and energy source to the yeast, as well as aspirating systems providing vigorous aeration
organic nitrogen, minerals, sulfur, vitamins, and to maintain highly aerobic growth. In the fer-
trace elements. If available, beet molasses is menter wort, 46% of yeast dry solids can be
blended with 1530% of cane molasses as a produced in a 15 h process.
vitamin (biotin) source. Additional nitrogen Postfermentation processing begins with
(ammonia or ammonium salts) and phosphate, separation of the yeast from the wort by centri-
minerals (calcium and magnesium), vitamins, fugation and counter-flow washing steps, to
and trace elements are needed, according to the produce a yeast cream with 1820% solids. The
composition of the yeast. If 100% cane molasses yeast cream is sold either as truck loads or
are used, thiamine and pantothenic acid have to containers to industrial bakers. The well-cooled
be supplied. Cane molasses is not acidified prior yeast cream stock in the yeast plant will be
to sterilization. press- or vacuum-filtered to a semisolid yeast
Table 1 shows the specific consumption mass of 30% solids. The filter cake is packaged
figures of bakers yeast production. as crumbly, bulk granular yeast in 25 kg poly-
Yeast is grown by the fed-batch process ethylene bags or extruded into 1 kg compressed
after several stages from pure culture to yeast blocks that are wrapped in wax paper or
seed I, seed II, and the commercial yeast stage composite films. It is cooled and shipped
in large fermenters of about 200 m3 work refrigerated to bakeries.
volume. The fermenters are equipped with cool- Instant active dry yeast (IADY) is produced
ing circle loops with plate heat exchangers to from the yeast filter cake by extrusion with
remove the excess heat energy, and jet spargers emulsifier agent as thin cylindrical strands.
These are dried on semicontinuous, two-stage
fluid-bed dryers. IADY with a solid content
Table 1. Specific consumption of raw material, process aids, and of 9296% is shipped in vacuum-packed or
energy in fresh bakers yeast (27% dry solid) and IADY (96% dry
solid) production nitrogen-flushed pouches. It does not require
refrigerated shipment or storage.
Specific consumption, t Wastewater from molasses conditioning,
fermentation, and washing steps need to be
Fresh bakers IADY
yeast treated. The specific organic load per ton of
molasses ranges from 180 to 210 kg COD-O2
Beet molasses (50% sugar), t 1.081.12 4.14.3
Ammonia (strong water), kg 23 80
or 140 to 170 kg BOD5-O2, respectively. The
Phosphoric acid (85%), kg 13 46 concentration depends on the overall process
Sulfuric acid (96%)a, kg 14 50 water use. The organic load can be degraded to
Magnesium sulfate  7 H2O, kg 2 7 biogas, if the BOD5-O2 concentration is over
Zinc sulfate  7 H2O, g 90 320
Biotin (2%), g 7 25
2 kg/m3. The degradation of COD-O2 from
Antifoam agent, L 0.5 2 nonassimilable molasses ingredients is more
Potato starch (filter aid, pre-coat), 12 50b complex. If flash evaporation is applied, this
kg matter and the salts can be concentrated and
Salt, kg 18 65b
Sodium carbonate, kg 814
sold as vinasse.
Extrusion aid, L 0.31.5
Emulsifier, kg 18 Yeast Strains. Strains of bakers yeast for
Cleaning and disinfection aid, L 1.2 5 industrial production are preserved as pure cul-
Process water, m3 1418 4365
Cooling water (< 15 C)c, m3 75 320
tures, usually on agar slants in test tubes. They
Steam (8 bar), kg 430 2950 are frequently transferred by well-known pure-
Electric power, kWh 285 1990d culture methods. Commercial fermentations are
Wastewater capacity, m3 1.42.0 57 started from such slants. Industrial strains of
Depending on the buffer capacity of molasses and the use of bakers yeast are quite stable. Bakers yeast
cane molasses (> 15%) strains are available from public and private
Only for rotary vacuum filter and cytorrhysis application, not
for chamber press filter
culture collections, and they can also be isolated
Depending on the climate zone and the process air conditioning from commercial bakers yeast, since this
Depending on the drying technology yeast enters the market as a viable product.
Yeasts 7

There are several patents protecting specific between 50 and 55 wt%. Molasses can be stored
strains. Enforcement of such patent rights well in large tanks often holding several thou-
depends, of course, on the ability to identify sand tons. It can be pumped at temperatures
the patented strain with certainty, which is often above 20 C. Molasses contains some insoluble
difficult. materials and some precipitation takes place on
Additional strains have been developed be- heating. Therefore, molasses is clarified. Cane
cause of the requirements of the baking indus- molasses cannot be filtered well, due to colloidal
try: a dry yeast strain with improved fermenta- substances present. It is clarified by addition of
tion activity, a yeast strain with exceptional sulfuric acid and sterilization followed by cen-
fermentation activity in high-sugar dough, and trifugation. Beet molasses can be clarified in the
yeast with improved performance in yeast-leav- same manner or by filtration. Prior to clarifica-
ened frozen dough. Some new strains have tion, molasses is diluted with water to 3040
become available, but acceptance by the baking Brix, which facilitates the process.
industry has been slow, partly because bakers do Some molasses components are not satisfac-
not like to deal with several strains in their tory for the production of yeast. The presence of
bakeries, and partly because the production and yeast-growth inhibitors, which may include
distribution of several strains by yeast compa- smaller fatty acids, nitrites, sulfites, and various
nies are more costly. Nevertheless, a trend insecticides or herbicides used in the field, or
towards more yeast strains for specialty bactericides, antifoam agents, and antiscaling
applications can be expected. surfactants used in the sugar factory.
Bakers yeast claimed organic may be Beet molasses contains 11.5 wt% raffinose
derived from the same strains as conventional on dry solid basis. Bakers yeast strains assimi-
bakers yeast but need to be grown on the late only the fructose moiety of the raffinose
organic sources for carbon and nitrogen and molecule, which accounts for one-third of the
complex sources for all the other supplements, trisaccharide content.
as no fertilizer-like substances are allowed Nitrogen. Beet or cane molasses contains
according to the standards (EC No. 1245/2008). low concentrations of nitrogenous materials.
Betaine nitrogen in beet molasses is not assimi-
Raw Materials. Carbon and Energy lated. A minor fraction of the low amino acid
Sources. Historically the major source of carbon content of molasses is assimilated (nitrogen
for yeast growth was maltose derived from malt- assimilation factor, NAF). In the production of
converted grain mashes. Today, glucose, fruc- bakers yeast, nitrogen is supplied in a readily
tose, sucrose, and a portion of raffinose from assimilable form as ammonia, ammonium phos-
molasses serve universally as the carbon source. phate, ammonium sulfate, or urea.
The discussion here is restricted to the growth of Feeding of nitrogen in the form of ammoni-
S. cerevisiae on molasses and a brief indication um sulfate or phosphate leads to liberation of
of potential sugar sources from starches and sulfuric acid or phosphoric acid, which must be
whey. neutralized to maintain the desired pH range.
Cane molasses is generally used in subtropi- When urea is used as a nitrogen source, fermen-
cal areas where cane is grown or in places to tation requires higher levels of biotin and
which it can be readily shipped by sea. In magnesium as cofactors.
northern climates where sugar beet is extensive- Minerals. Due to the low level of phosphorus
ly grown, and in areas distant from marine in molasses, additional P should be added to the
transportation, beet molasses is the major con- growth medium. The required concentration of
stituent of the mash. The price of molasses is phosphorus expressed as P2O5 should be one-
largely determined by its value as a feed. In third of the nitrogen concentration. Molasses
fermentation industry one of its major uses is the contains sufficient potassium, calcium, and
production of bakers yeast, followed by ethanol sulfur, but is low in magnesium, which must
biofuel production. At present, molasses is the be added to the growth medium.
cheapest source of fermentable sugars. Vitamins. Bakers yeast strains require biotin
Beet and cane molasses are shipped at 8085 for growth. Cane molasses supplies ample
Brix. They have a sugar content that varies amounts of this vitamin, usually 0.50.8 ppm.
8 Yeasts

Beet molasses contains only traces of biotin. Specifications for bakers yeast, like the
Therefore, at least 15% cane molasses is used in General characteristics of fresh/dry bakers
beet/cane blends for yeast production. For fer- yeast, published by the European association
mentation of carbohydrate sources that contain of yeast producers, COFALEC, require absence
little or no biotin (e.g., corn syrup and date of Salmonella and other pathogens, and restrict
concentrate), the synthetic vitamin can be added the number of coliform organisms to less than
at a rate of 60100 mg per 100 g of yeast solids 1000 per gram and the count of Escherichia coli
grown. to less than 100 per gram. Bakers yeast plants in
Beet molasses contains sufficient amounts of the US follow Good Manufacturing Practice
other vitamins including pantothenate, thiamin, (GMP) rules of the Food and Drug Administra-
pyridoxine, and nicotinic acid. Bakers yeast tion (FDA), and great efforts are made to keep
can synthesize thiamin from thiazole and the number of contaminants to a minimum.
pyrimidine compounds, and there is no absolute Molasses is generally sterilized in bulk
need for thiamin, but there is a demonstrable or in continuous heat exchangers. The air
effect on the yeast fermentation activity. used to aerate the fermenter is passed through
Alternative Carbon Substrates. Molasses is depth filters or membrane filters, or is heated.
the least expensive source of sugar for growing Fermentation tanks are thoroughly cleaned by
bakers yeast, but its price occasionally fluctu- hot alkaline solutions or detergents, usually
ates and may approach that of corn-derived with clean-in-place (CIP) equipment, and if
sugars. Therefore, other carbohydrate sources necessary by scrubbing. Desinfectants are
have been investigated, including hydrolyzed quartenary ammonia salts or peroxoacetic
lactose from cheese whey, and hydrolyzed acid.
starch. For growth of bakers yeast the hydroly- All pipes, pumps, valves, and other auxiliary
sis of starch by commercial amylases would be equipment must be thoroughly sanitized. It is
more cost effective. also important to consider potential contamina-
If sugar syrups from sugar mills or any tion from other accessory equipment such as air
hydrolyzed starch source are used, the medium pipes, leaking cooling circle heat exchangers,
has to be supplemented with potassium salts and probes for pH meters and other instruments and
others, relevant for the osmotic pressure condi- sampling devices. Particular attention has to be
tions in the wort. paid to ducts for exit air to prevent condensate
Fermentation Activators and Inhibitors [4]. from dripping back into the fermenter. Similar
Activators are the nutrients required for the attention must be paid to further downstream
successful completion of the fermentation pro- processing of the grown yeast, which includes
cess. Activators include P, Mg, K, Na, Mn, and the centrifuges, presses, filters, and all packag-
S. Only the trace elements (Cu, Fe, and Zn) are ing operations. For ADY it includes the dryers
essential for yeast metabolism. They become as well. Yeast is an excellent growth medium for
integral part of the proteins as activators of bacterial contaminants, and deposits of wet or
enzymes and function as cofactors in biochemi- dry yeasts on equipment become a ready source
cal reactions within the cell. of infection.
Fermentation inhibitors include heavy me- The overwhelming proportion of contami-
tals at very low concentrations. SO2 also inhibits nants in CY or ADY are lactic acid producing
yeast growth, but there is good adaptation even bacteria (LAB), which almost always belong to
at concentrations of 80100 ppm. Molasses the heterofermentive lactic acid producing bac-
contains variable amounts of nitrate that can be teria of the genus Leuconostoc or to the homo-
reduced to nitrite by bacterial action during fermentive bacteria of the genus Lactobacillus.
growth. Yield losses due to nitrite inhibition Some coliform organisms and occasionally a
have been reported for nitrite concentrations of few E. coli can be found. ADY made by drying
0.0010.004%. CY contains the same kinds of microorganisms,
Contamination. Bakers yeast always con- but the total count generally decreases during
tains some contaminating organisms since it is drying and upon subsequent storage of the dried
grown in fermenters that cannot be kept yeast. Specifications for rope spores require an
completely sterile. upper limit of 200 per gram ADY.
Yeasts 9

Occasionally contamination with Odium lac- (IADY) because it does not require separate
tis (milk mold) or other molds can occur. Vari- rehydration before use.
ous species of contaminating wild yeast have Numerous patents have claimed certain
also been reported, most of them falling into the strains of S. cerevisiae as particularly useful in
genera Candida and Torulopsis. the production of IADY, often in conjunction
with definite processing conditions. Such strains
are produced by classical hybridization, proto-
3.2.2. Active Dry Yeast (ADY) and Instant plast fusion, or mutation.
Active Dry Yeast (IADY) [3] Many compounds added to the yeast cream
or to the press cake are claimed to produce a
Bakers yeast can be dried in its vegetative more active ADY or IADY. Emulsifiers such as
state to a product having less than 8% monoglycerides, soya lecithin, glycerol polye-
moisture, without substantial loss of viability, sters, and sorbitan esters also help maintain the
but success of the drying method requires yeast activity. In practice, sorbitan esters are
mastery of the various requirements of the widely used at levels of 0.21%. The added
methodology. emulsifier facilitates processing and lightens the
For a given strain, the fermentation activity is color of ADY or IADY, but the main reason for
considerably higher in yeast grown to a higher the use of these compounds is the reduction of
nitrogen level, but such yeasts are also more leached solids when ADY is rehydrated.
sensitive to drying. With regard to composition, Active dry yeast loses activity upon storage
it was found that the trehalose concentration when exposed to atmospheric oxygen. Storage
should be 12% or higher (solid basis). It was also stability can be extended by complete exclusion
found that compressed yeast used for the pro- of air during storage.
duction of ADY must be thoroughly washed to For use in baking ADY must be rehydrated in
reduce the concentration of water-soluble sub- water at 3040 C. Rehydration at lower water
stances, especially residual salts that are partic- temperatures lessens the fermentation activity
ularly harmful during drying. of ADY, due to the extraction of reducing
American Type Culture Collection (ATCC) compounds such as glutathione or cysteine,
ADY strain no. 7752 was grown to a nitrogen which leads to slackening of the dough. Slack-
level rarely exceeding 7% of dry solid. The ening may be desired for industrial machined
lower nitrogen level favored the stability of the baking or when very strong, gluten-rich flours
yeast but resulted in a somewhat lower fermen- are applied, but it is better achieved in a con-
tation activity in comparison with that of com- trolled fashion by the addition of proteolytic
pressed yeast (nitrogen 8.28.8% of dry solid) enzymes or cysteine to the dough.
on an equivalent solid basis. There is some loss IADY can be mixed in with other dry dough
of solids during the early stages of drying be- ingredients directly and needs no prior rehydra-
cause of the increased endogenous respiration tion in water. Its activity is restored by the flour
and assimilation of the storage carbohydrates. moisture content.
This strain can be dried in continuous tunnel The storage stability of ADY and IADY
(chamber) dryers with a minimal loss of decreases at higher temperatures, higher mois-
fermentation activity. ture content of the yeast, and by the presence
Hence, Gist Brocades developed the fluid- of air in the package. For storage under
bed drying method, which employs drying at a nitrogen or when vacuum-packed, the loss is
lower temperature in a shorter period of time about 1% per month and generally less than
and with much smaller particles of press cake, 10% per year.
which improves the uniformity of moisture Accelerated tests, e.g., at 43  C for one or two
removal. Their patent comprised a drying time weeks, are useful to simulate yeast quality
of < 2 h and a yeast strain with a nitrogen changes during commercial long-term storage
content of more than 8%. The resulting ADY at normal temperatures after one or two
has a smaller particle size and more regular years. Accelerated storage tests under the effect
shape than ADY dried in tunnel dryers. This of various atmospheres and temperatures
type of dry yeast is called instant active dry yeast approaching 55 C have been conducted.
10 Yeasts

ADY readily absorbs CO2. When used with 3.2.4. Dairy Yeasts
large packages, a sufficient vacuum is created to
give the appearance and performance of a vac- Dairy yeasts are lactose-assimilating and lac-
uum packaging. Once the package has been tose-fermenting yeasts; the preferred species for
opened the ADY should be used quickly. In the production of biomass is Kluyveromyces
particular, it should not be exposed to an atmo- marxianus.
sphere with high humidity. Large amounts of cheese whey are
IADY, which is even more hygroscopic, is available as a byproduct of cheese production
always distributed in hermetically sealed, nitro- (! Cheese, Processed Cheese, and Whey). The
gen-flushed aluminum foil pouches, with low whey may be dried and sold as food or as feed
permeability to water vapor and gaseous ingredients, but supply by far surpasses demand.
oxygen. The aluminum foil is laminated to a Thus, there exists a problem of waste disposal
heat-sealable plastic film. and there have been numerous attempts to use
whey as a fermentation substrate for the
production of ethanol and biomass. One ap-
3.2.3. Candida utilis (Torula) Yeast proach is to digest the whey sugars by enzymes
to galactose, which can be assimilated by most
Torula is a common commercial trade name of the Saccharomyces strains as well.
for dried nutritional and feed yeasts. The
major substrate for its production in the United
States is sulfite waste liquors of the paper pulp 3.2.5. Brewers Yeast
process, and in Europe and former Eastern Bloc
countries the substrate is wood hydrolysates or Yeast plays an important role in the brewing
sulfite waste. C. utilis is grown on wood sugars process not only by converting fermentable
because it assimilates both pentoses and sugars in the wort to ethanol and carbon dioxide
hexoses, and its growth requires the addition but also by producing a variety of volatile and
of nitrogen sources, phosphate, and potassium nonvolatile constituents, mostly in trace
to the medium. In contrast to S. cerevisiae, amounts, that contribute to the whole flavor of
Candida yeast does not require addition of biotin the beer.
for growth. Two species of the genus Saccharomyces are
C. utilis is produced by continuous fermen- often considered important for brewing, de-
tation, with high aeration in specially designed pending on the type of the brewing process. For
foam fermenters capable of 60% air content. more details on the brewing processes, see
Providing high-quality inoculum, fermentation ! Beer.
can run for months. The yield of yeast is gener- Most beers are produced by the bottom-fer-
ally 50% based on fermentable pentoses plus mentation process in which strains of the yeast
hexoses. The yeast is recovered by centrifuga- S. uvarum are used. These yeasts strains tend to
tion. After minimal washing with water, C. utilis remain suspended for a limited time in the
can be sold as feed yeast, which contains a fermentation medium while the fermentation is
considerable amount of lignosulfonic acids. For active, after which they settle to the bottom of
use as human food, the yeast must be centri- the fermentation vessel. These strains are pre-
fuged at least twice with extensive washing. dominantly used in Germany and North Amer-
Inactive dried nutritional yeast is produced ica for the production of lager beer.
from a yeast cream of about 18% solids by The other type of beer, referred to as ale, is
pasteurization followed by drum drying or spray produced by a top-fermenting yeast such as
drying. The spray-dried product has very fine S. cerevisiae. The strains tend to rise to the top
particles. The drum-dried product exiting the of the fermenting medium and are generally
drums as a thin sheet is available as small flakes skimmed off the surface of the fermenting liquid
or ground to a powder. In countries that lack after the fermentation is completed. Ales are
cheap sources of oil-seed meals (mainly soy- popular in England and Ireland.
bean meal), C. utilis biomass is used extensively Malt extract or beer wort is one of the oldest
as a protein supplement. classical media for growing yeast cultures. It
Yeasts 11

contains nitrogen, mineral salts, and vitamins in readily transferred during the crushing season
amounts adequate for maintaining yeast growth, by fruit flies (Drosophila melanogaster) during
and fermentable carbohydrates to supply carbon the transport of grapes to the winery and within
and energy for biosynthesis. In brewers yeast the winery. Once the crushing season is under-
propagation, there is also an absolute require- way, the major source of natural yeasts is likely
ment for molecular oxygen, provided by small- to be found on winery equipment such as
inline jet aeration. The C-to-N ratio of the wort crushers, destemmers, and tanks.
can be improved by using malt sprouts as addi- There is a considerable increase in cell
tional nitrogen source. The yeast can, however, counts as the grapes are transported and the
exhibit some growth under completely anaero- must is prepared in the winery. The fermenta-
bic conditions. tion is initiated by weakly fermenting yeast,
Spent excess brewers yeast is recovered in mainly Kloeckera apiculata. For reasons of
breweries by sedimentation or centrifugation. It fermentation efficiency and reliable flavor for-
contains 1216% total solids, of which approx- mation, the early fermenters are replaced by
imately two-thirds are yeast solids and one-third more alcohol-tolerant yeasts, and during the
are beer solids. The slurry must be kept refrig- main part of the fermentation and toward the
erated before use because even at 4 C, respira- end the true wine yeasts S. cerevisiae dominate.
tion lowers the carbohydrate content. This yeast
may be pasteurized and dried as such to a rather
3.2.7. Distillers Yeasts
bitter dried brewers yeast, or the yeast may be
separated from the beer by decanter or nozzle
For more detailed information on the production
centrifuges and partially debittered. It is also
of spirits, see ! Spirits.
pasteurized and dried. Yeast biomass in its
Distillers yeasts include strains of S.
inactive dried form is used widely as feed
cerevisiae, except for natural fermentation, in
supplement. It is used in poultry rations and pig
which any number of species may be involved.
starter feeds. The use of brewers and distillers
The fermentation of the various distilling raw
byproduct yeast in feed has been demonstrated.
materials need to be categorized into the fer-
mentation of cereal mashes (whisky and sake)
and the natural wine fermentation of juices
3.2.6. Wine Yeasts
and liquid extracts, which contain all carbohy-
drates already in the form of fermentable sugars
For details on wine production, see ! Wine.
and fully hydrolyzed starches.
Fermentations of fruits and fruit juices occur
Cereal mashes undergo the concurrent enzy-
naturally and spontaneously without human
matic formation of sugars, either prior to or
intervention. Ripe berries can undergo sponta-
simultaneously to the fermentation process.
neous fermentation in the field if they are over-
Yeasts suitable for cereal mash fermentations
ripe or damaged. These fermentations are initi-
can be bakers compressed yeast, distillers
ated by yeasts found on the grape skin. In a
active dry yeast, or strains grown in the distillery
broad sense, any yeast species occurring on
from selected pure-culture slants.
grapes and participating in must fermentation,
Distillers yeast is not recovered as a separate
no matter how transient, may be called a natural
product because of the presence of other inso-
wine yeast. In a narrow sense, only strains
lubles in the still residues and is sold as
grown on pure culture in winery fermentations
distillers dried grains with solubles (DDGS)
are wine yeasts. Such strains generally belong to
protein-rich product to the feed industry.
the species S. cerevisiae and provide a variety of
flavors to build a certain wine taste.
A large number of yeast cells found on grapes 4. Yeast-Derived Products
belong to the genera Kloeckera, Hanseniaspora,
and to a lesser extent to the genera Metschni- 4.1. Flavor Products
kowia and Candida.
Natural yeasts survive from year to year in Yeast extracts comprise proteins, free
the intestines of bees and wasps and they are amino acids, vitamins, and minerals and are
12 Yeasts

commercially available as a powder or concen- blood plasma levels of uric acid and may
trated paste with 60% solids. Surplus yeast from cause gout. Decrease of nucleic acid content
the brewing industry is relatively cheap and is is often carried out following the separation of
utilized in the production of yeast extracts to yeast proteins in crude form from the whole
meet the needs of the food and fermentation cell. Yeast is ruptured using several methods
industries. including high-pressure homogenization, col-
The yeast extracts are generally made by loid mill, sonic disintegration, freezethaw
autolysis, which employs the yeasts own pro- treatment, use of lytic enzymes, or a combi-
teolytic enzymes to solubilize the within-cell nation of these methods for maximum recov-
components at elevated temperatures of >45 C. ery. Several methods to lower the level of
Under these conditions, the cell walls or cell nucleic acids have been described in the liter-
hulls remain intact and are separated as a ature. There is no need, however, for the
byproduct by centrifugation. removal of nucleic acids from yeast biomass
Yeast extracts can also be made by hydro- for use in feed.
lysis with external reagents or enzymes
capable of releasing the cell contents. Extracts
made from primary-grown yeasts such as 4.3. Colorants Derived from Yeast
S. cerevisiae (bakers yeast), C. utilis, or
K. marxianus have flavor profiles somewhat There is considerable interest in developing
different from those of brewers yeast. The food colorants from natural products to replace
latter is likely to have undesirable flavor the synthetic ones, especially in the diet of
characteristics as a result of a carryover of children. Although different types of natural
hop resins and beer solids. pigments are widely distributed in the plant and
Yeast extracts have received wide accep- animal kingdoms, a variety of carotenoid pig-
tance in the food industry as natural flavors. ments of various pink and yellow shades are also
They are also cost-effective compared to other produced by a number of yeast species. Yeast
synthetic food flavors, on the basis of equivalent offers considerable advantages over other natu-
flavor intensity. Yeast extracts are used ral sources in that it is inexpensive to grow on an
extensively in the fermentation industry as a industrial scale. Yeast has now begun to emerge
complex substrate and in analytics for the prep- as a cost-effective source for the production of
aration of microbiological growth media for natural food colorants.
plate assays. The best-known genera among the pigmen-
Autolysate is the total content of yeast fol- ted yeasts are Rhodotorula, Rhodospondium,
lowing the autolysis degradation process. The Cryptococcus, Sporidiobolus, and Sporobolo-
product is in the form of a concentrated paste or myces. These yeasts are characterized by their
dried, without separation of the cell-wall matter. ability to produce pigments such as b-carotene,
Consequently, autolysates are less intense in a-carotene, torulene, torularhodin, plectanis-
flavor than extracts on an equal weight basis xanthin, and 2-hydroxyplectanisxanthin.
because of the dilution effect of the cell-wall Carotenoids produced by different Rhodo-
material. torula species primarily contain the two oxy-
genated compounds torulene and torularhodin,
and the ratio of the two major pigments deter-
4.2. Nutritional Yeast mines the intensity of the shades of red in
different species.
Yeast is generally recognized as an excellent Phaffia rhodozyma is a basidiomycetous
source of proteins, B vitamins, fiber, and many yeast that differs from other pigmented yeasts
other micronutrients. However, for the use as a in producing astaxanthin, which is the charac-
major source of protein by humans the teristic pink color in the plumage of birds such
presence of nucleic acids is a serious obstacle, as flamingos, in the exoskeleton of marine in-
and nucleic acid nitrogen accounts for about vertebrates such as lobsters, crabs, and shrimps.
1015% of the total nitrogen of yeast cells. In fish like trout and salmon, the pigment is
The intake of nucleic acids leads to elevated stored in their flesh. Lack of the characteristic
Yeasts 13

pink color in pen-reared salmonids renders This enzyme is widely used in the food
them less acceptable. Astaxanthin can restore industry for the treatment of frozen whole-milk
the pink color in seafood, resulting in better concentrates to prevent crystallization during
marketability. storage, and most importantly to make milk-
based products acceptable for those who cannot
tolerate lactose.
4.4. Yeast-Derived Enzymes [4] a-Galactosidase hydrolyzes the disaccha-
ride melibiose to glucose and galactose as well
Invertase is an enzyme capable of splitting as the trisaccharide raffinose to sucrose and
the sucrose molecule into glucose and fructose, galactose and can improve the production yield
a noncrystallizable mixture known as invert in the beet sugar industry. a-Galactosidase is
sugar. Bakers yeast is rich in invertase and is not present in bakers yeast, but in brewers
the main source of commercially available yeast (S. uvarum).
invertase. A potential application for a-galactosidase is
The most common method of extracting and in the bakers yeast industry. Although the
concentrating invertase from yeast is by autoly- predominant sugar in both beet and cane molas-
sis and to recover the enzyme from the filtered ses is sucrose, beet molasses contains 11.5%
autolysate by precipitation or concentration. raffinose (solid basis). Only one-third of this
The precipitating agents commonly used are trisaccharide is utilized by bakers yeast due to
ethanol, acetone, and ammonium sulfate. The the presence of invertase, which converts raffi-
precipitate thus formed is dissolved in water, nose into melibiose and fructose. The latter is
filtered, and treated with activated carbon to assimilated by the yeast. For complete utiliza-
remove coloring matter, followed by concentra- tion of raffinose molecule the enzyme a-galac-
tion under vacuum, or by ultrafiltration. Inver- tosidase can be expressed in a genome-modified
tase is commercially available as a concentrate bakers yeast.
stabilized with glycerol. Yeast-Derived Biologically Active Com-
There are two important applications of pounds. From different yeasts, several enzymes
invertase in the food industry. The first is in and biologically active compounds are produced
the manufacturing of confectioneries, where on a commercial scale, such as alcohol dehydro-
invertase is used mainly in the production genase; hexokinase; L-lactate dehydrogenase;
of soft-centered candies (! Confectionery). glucose-6-phosphate dehydrogenase; glyceral-
The use of invertase provides the plastic dehyde-3-phosphate dehydrogenase; inorganic
consistency to the center and prevents sucrose phosphates; coenzyme-A; oxidized and reduced
from crystallizing during storage. The enzy- diphosphopyridine nucleotides; and nucleotides
matic hydrolysis that brings about the creamy like mono-, di-, and triphosphates of adenine,
consistency of the center generally takes place cytidine, guanosine, and uridine.
after casting and covering the hard center with
chocolate. The second important application
is in the inversion of syrups for the production 4.5. Cosmetic and Pharmaceutical
of high-test molasses, used extensively in the Products [1]
fermentation industry. High-test molasses, if
not inverted, would soon tend to crystallize Skin Respiratory Factor. A live-yeast-cell
because of its high concentration of sucrose. derivative (LYCD) product, comprising several
Inversion also raises the osmotic pressure stress metabolites of yeast, is often referred to as
and results in a microbiologically stable con- skin respiratory factor (SRF). SRF prepared
centrate with a consistency that is easy to from yeast is known to exhibit certain biological
handle. properties capable of enhancing the oxygen
Lactase catalyzes the hydrolysis of lactose uptake rate of tissue, thereby stimulating wound
(milk sugar) into its principal components, healing. Its effectiveness in skin care and ageing
glucose and galactose. Lactase is not present in protection has made it a commercially attractive
bakers yeast, but it is present in a number of ingredient for the cosmetic and pharmaceutical
other common yeasts such as K. marxianus. industries.
14 Yeasts

Glucose Tolerance Factor. Diets supple- nutritionally nonfunctional in the human diges-
mented with dry yeast, comprising trace tive tract. The three components account for
elements such as chromium, selenium, and more than 80% of the dry matter of the glycan
molybdenum, enable a partially or wholly fraction, and about 20% is noncarbohydrate
alleviation of diabetes type II problems. material, including protein, nucleic acid, lipids,
An active component influenced by such and ash. Therefore, a desirable property of
diets is a trivalent chromium complex, referred glycan is its ability to function as a noncaloric
to as glucose tolerance factor (GTF). Patients in food thickener
the diabetes type II group have GTF early in life, More recent studies have demonstrated the
but tend to lose it with age, causing increased beneficial effect of glycan in improving host
vulnerability to diabetes. It is suggested that resistance by enhancing both humoral and cel-
GTF functions as a cofactor for insulin, thereby lular immunity to certain malignant tumors and
enhancing the binding of insulin to receptive viral infections. The therapeutic effect of glycan
sites on the membranes of insulin-sensitive is primarily due to the stimulation of the reticu-
tissues. loendothelial system (RES), which in turn pro-
Other investigations showed a reduction in duces increased amounts of macrophages that
the level of cholesterol and triglycerides in the play a key role in the bodys natural immune
blood of humans supplementing their diets system. The macrophages absorb and destroy
with brewers yeast rich in GTF. However, invading particles by phagocytosis.
further feeding studies are necessary to confirm
the results before yeasts rich in GTF could be
recommended as a possible treatment for 4.6. Yeast Engineering [5]
lowering blood sugar or curing lipid disorders.
With the technological advances that have
Selenium Yeast. Selenium has been recog- already been made in genetic engineering,
nized as an essential trace element for both substantial effort has now gone into the devel-
human and animal nutrition, with regard to both opment of genomics and metabolomics meth-
cardiovascular fitness and cancer prevention. odology to manipulate and modify the yeast
Selenium yeasts comprise selenomethionine genome and metabolic flux. Yeast has become
and selenocystein as active ingredients. Organ- one of the most frequently exploited host micro-
ically bound Se, such as selenomethionine, is organisms, capable of producing valuable traits
incorporated several times faster into the body of commercial interest:
tissue than inorganic Se.
Surveys in some parts of Finland have shown . In frozen food (dough) application, where,
that people with a blood selenium level lower e.g., LT clones of yeasts were constructed to
than 0.04 mg/L are three times more prone to enable silencing the bakers yeast raising
heart attacks than their counterparts with normal power until a certain temperature is reached.
selenium levels. . In bioethanol application, where cellulase
Organically bound Se is now recognized from basidomycetes can be expressed in ge-
as nutritionally important because of its nome-modified yeasts to enable fermentation
ability to prevent some vitamin E deficiency of cellulosic material from grass, straw, and
disorders, at least in laboratory animals fed diets corn stover.
partially reduced in nucleic acids. Nutritional . In white biotechnology, where, e.g., metabol-
yeasts with an intracellular Se concentration of ic engineering make the yeast cells accumu-
1000 ppm are commercially available as dietary lating C4 metabolites as building-block
supplements. chemicals.

Glycan. The term glycan is used broadly Despite the exceedingly long lag time and
to refer to a yeast cell-wall byproduct during the high expense of putting new products through
production of soluble extracts from bakers or the entire risk study and approval process, at
brewers yeast. The yeast cell wall carbohydrate least a few biotechnology companies have been
polymers glucan, mannan, and chitin are able to bring certain genetically engineered
Yeasts 15

yeast-derived products into the final stages of 2 A.H. Rose, J.S. Harrison: Yeast Technology in The Yeast,
marketing. vol. 5, Academic Press, London 1993.
3 K. Fischer, J. Rahn, W.K. Bronn: Hefe und Hefeextrakte in
R. Heiss (ed.): Lebensmitteltechnologie, 6th ed., Springer, Berlin
2004, chap. 40.
4 G.M. Walker: Yeast Physiology and Biotechnology, John Wiley
References and Sons, Hoboken, NJ, 1998.
5 H. Feldmann: Yeast, Wiley-VCH, Weinheim 2010.
1 G. Reed, T.W. Nagodawithana: Yeast Technology, 2nd ed., AVI,
New York 1991.