Shurson 2018 Yeast and Yeast

Animal Feed Science and Technology 235 (2018) 60–76
Contents lists available at ScienceDirect
Animal Feed Science and Technology

journal homepage: www.elsevier.com/locate/anifeedsci
Review Article
Yeast and yeast derivatives in feed additives and ingredients:

T
Sources, characteristics, animal responses, and quantification
methods
⁎
G.C. Shurson
Department of Animal Science, 385 ANSC/VM Bldg., 1988 Fitch Ave., University of Minnesota, St. Paul, MN 55108, USA
AR TI CLE I NF O AB S T R A CT
Keywords: Numerous yeast products and yeast-containing feed ingredients are commercially produced,
Yeast marketed, and used extensively in animal feeds around the world. Considerable research has
Feed additives been conducted to evaluate the potential animal growth performance and health benefits of
Distillers co-products adding yeast, yeast-derivatives, and yeast-containing ingredients into animal feeds. Active dry
Immune responses
yeasts are commonly used solely or in combination with beneficial bacteria in probiotic products.
Growth performance
Nutritional yeasts are used as supplements in animal feeds due to their relatively high protein and
Analytical methods
amino acid, energy, and micronutrient content compared with common feed grains and oilseed
meals. Other important yeast-based products contain nutraceutical compounds present in yeast
cells and cell walls (i.e. β-glucans, mannanoligosaccharides, nucleotides) that have generally
been shown to improve animal growth performance and health. Specialty yeast products, such as
selenium yeast (highly concentrated and bioavailable source of selenium) and Phaffia rhodozyma
yeast (contains pigment that improves flesh color in salmon and trout) have specific applications
in some animal feeds. Ethanol co-products such as corn distillers dried grains with solubles
(DDGS) and new grains distillers dried yeast ingredients, containing more than 40% crude
protein, also contain significant amounts of yeast cell and nutraceutical components. Therefore,
because these yeast-based products have several nutritional and health benefits, they are be-
coming alternative supplements in animal feed due to restrictions on antimicrobial growth
promoter use in many countries. However, it is difficult for nutritionists to differentiate the
characteristics, composition, and optimal feeding applications among the diverse number of
yeast-containing products available. Furthermore, most of these products contain combinations
of probiotics and nutraceutical compounds with different modes of action, making it difficult to
determine which compounds contribute to specific responses observed. Quantification of these
nutraceutical compounds is difficult, and except for methods to determine viable yeast in dried
active yeast products, there are no standard methods for determining dead yeast concentration or
fast, inexpensive, and accurate methods to estimate the proportion of yeast components in var-
ious yeast-containing additives and feed ingredients. Due to the increasing popularity of using
yeast-based products in animal feeds, development of analytical approaches to estimate yeast and
its components in these products is greatly needed. In this review, various categories of com-
mercially available yeast and yeast-containing additives and feed ingredients will be described
along with our current knowledge about their role in improving animal growth performance,
Abbreviations: AAFCO, Association of American Feed Control Officials; ADG, average daily gain; ADFI, average daily feed intake; CFU, colony forming units; DDGS,
distillers dried grains with solubles; DFM, direct fed microbial; DNA, deoxyribonucleic acid; EPA, Environmental Protection Agency; IFN, international feed names;
MOS, mannanoligosaccharides; N/A, not applicable; RNA, ribonucleic acid
⁎
Corresponding author at: Department of Animal Science, 335d ANSC/VM Bldg., 1988 Fitch Ave., University of Minnesota, St. Paul, MN 55108, USA.
E-mail address: [email protected].
https://doi.org/10.1016/j.anifeedsci.2017.11.010
Received 30 June 2017; Received in revised form 14 November 2017; Accepted 15 November 2017
0377-8401/ © 2017 Elsevier B.V. All rights reserved.
G.C. Shurson Animal Feed Science and Technology 235 (2018) 60–76
health, and proposed mechanisms of action, and challenges of quantifying yeast content and their
biologically active components.
1. Introduction
Yeasts are single cell, eukaryotic microorganisms classified in the fungi kingdom (Bennett, 1998; Ingraham, 2010). These mi-
croscopic fungi are generally about 3–4 μm in size, have a nuclear membrane and cell walls, but unlike plants, they contain no
chloroplasts. Yeasts are characterized as heterotrophs in which they rely on living and dead organic material as sources of energy and
nutrients (Bennett, 1998). Yeast cells obtain their nutrition by producing and releasing various proteolytic, glycolytic, or lipolytic
enzymes to digest organic matter, or by absorbing amino acids and monosaccharides through the cell wall (Baron, 1996). Re-
production occurs by budding and fission. Budding occurs when a parent cell increases in size, and a protrusion forms along the cell
wall to form a “bud”, which breaks from the parent cell or is partially conjoined in elongated cells (Evans et al., 2000). Fission also
occurs when a parent cell divides into two daughter cells (Evans et al., 2000). Yeasts are considered facultative anaerobes which
means that they can survive and grow in the presence or absence of oxygen (Stone, 2006). Yeast propagation occurs under aerobic
conditions, and cells convert oxygen and sugars into carbon dioxide and energy through oxidative metabolism to allow efficient yeast
cell growth. Under anaerobic conditions, such as those used in beverage and fuel ethanol production, yeasts are much less efficient in
these processes, which results in the production of ethanol (Bekatorou et al., 2006).
Animals have been fed various forms of yeast and yeast derivatives for more than 100 years (Stone, 2006). However, recent
government restrictions and elimination of the use of growth promoting antibiotics in animal feed in the European Union and United
States have led to a significant increase in interest in using alternative products (including yeast products) to provide animal health
and growth performance benefits. Furthermore, feed ingredients produced from yeast fermentation processes (e.g. distillers dried
grains with solubles; DDGS) in animal feeds has increased dramatically in recent years (Shurson, 2017). Therefore, there are many
types of feed additives and feed ingredients that contain yeast in various forms, but animal nutritionists are often uncertain about the
unique differences among these products and their potential roles in animal nutrition and health. Furthermore, despite the fact that
yeast-containing additives and feed ingredients have been widely used in the feed industry for decades, there are no standard
analytical methods to quantify yeast and their biologically important chemical components. Unfortunately, limited information has
been published on the accuracy and use of practical methods to quantify yeast components for the feed industry. Our inability to
accurately quantify yeast and yeast components in feed additives and feed ingredients has become a significant issue requiring
investigation because of the need to accurately determine the dosage and diet inclusion rates to achieve desired concentrations of
biologically active substances. Quantification of yeast and yeast components is essential for achieving their desired potential benefits
in animal health and performance and preventing excessive feeding of biologically active components of yeast. Therefore, the
purpose of this review is to 1) describe the various forms of yeast in feed additives and feed ingredients, 2) briefly review their
benefits to animal health and growth performance, and 3) review various methods to quantify the concentrations of yeast and their
known active components.
2. Yeast species of commercial importance
Yeasts are found in abundant quantities and are almost everywhere in the environment. They have been isolated in fruit, honey,
soil, water, and plant stems, leaves, and flowers (U.S. EPA, 1997), and are naturally present in common feed ingredients such as
grains, grain co-products, silage, and hay fed to animals. Most species of yeast are neither harmful nor beneficial to humans and
animals. A few genera of yeast are known to be pathogenic (Candida, Cryptococcus, Torulopsis, and Trichosporon; Kandel and Stern,
1979), while some species (Saccharomyces cerevisiae, Kluyveromyces marxianus, Candida utilis) provide beneficial effects.
There are about 60 different genera of yeast, which are comprised of about 500 different species (Stone, 2006). Yeast species vary
in their cellular morphology, metabolism of different substrates, and reproduction processes (Stone, 2006). However, only a few of
these species are used commercially. Commercial applications for yeast include the production of alcoholic beverages (i.e. beer, wine,
and spirits), non-alcoholic beverages (i.e. root beer, kvass, kombucha, kefir, mauby), bread and pastry baking, bioremediation,
industrial ethanol production, nutritional supplements, probiotics, aquarium hobbies (to generate carbon dioxide for support plant
growth in aquaria), food additives and flavoring agents, scientific research, and genetically engineered biofactories. Saccharomyces
cerevisiae is the predominant species used in food, beverage (distilled spirits and beer), and fuel ethanol production processes, where
selected strains convert glucose and sucrose to ethanol (Reed and Nagodawithana, 1991). Other commercially important yeast strains
include Kluyveromyces marxianus (whey yeast) which uses glucose and lactose in milk as its substrates, and Candida utilis (Torula
yeast) which uses xylose and glucose in wood pulp from paper manufacturing as substrates (Reed and Nagodawithana, 1991).
The intracellular chemical components of yeast cells include amino acids, peptides, carbohydrates, salts, monosodium glutamate,
nucleic acids (RNA), enzymes, and cofactors (Hassan, 2011; Dubey et al., 2010). Yeast cell walls are comprised of glucans, glyco-
proteins, mannans, and chitin (Alexandre and Guilloux-Benatier, 2006; Kollar et al., 1997). The combination of these compounds
make them attractive not only as nutritional supplements in animal feeds, but also useful nutraceuticals.
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3. Yeast and yeast-containing feed ingredients
The official definitions and international feed names (IFN; if available) of the major yeast products and yeast containing feed
ingredients that are approved for use in animal feeds are available from AAFCO (2017).
3.1. Viable yeast products
Viable yeast products are generally added to animal feeds for their potential probiotic effects. The most common viable yeast used
in the animal feed industry is active dry yeast (AAFCO – 96.2 Active Dry Yeast; IFN 7-05-524 – Yeast active dehydrated), which
contains about 95% dry matter (Stone, 2006). Active dry yeast is also found in several direct-fed microbial feed additives (AAFCO –
36.14 Direct-Fed Microorganisms). Other forms of viable yeast include wet yeast cake (30% dry matter) and yeast cream (about 20%
dry matter), which are used extensively in the bakery industry (Stone, 2006). However, all three of these viable yeast forms can be
used as inoculum to manufacture yeast culture, which is a yeast fermented product also used in the feed industry (Stone, 2006).
Active dry yeast is comprised of 15–25 billion live yeast cells (colony forming units; CFU’s) per gram (Stone, 2006). The three
most common processing methods include tunnel dried yeast (granular powder), fluid-bed dried yeast (quick rise yeast in oval shaped
spheroids), and rotolouver dried yeast (produces small spheres or balls). The tunnel dried and fluid-bed drying methods are most
common in the U.S., while the rotolouver drying method is more common in Europe and Latin America (NPCS Board of Consultants
and Engineers, 2011). Of these drying processes, fluid-bed drying has become the most popular because it causes less damage to yeast
cells, and thus, maintains their viability.
In addition, pure, active dry yeast is also found in diluted feed additive products, which contain other ingredients such as rice
hulls and distillers solubles. Many of these diluted products contain only 5 × 109 CFU’s per gram, which represents 20–25% of the
CFU’s of pure, active dry yeast (Stone, 2006). As a result, it is important to compare the price of these products, on a cost per 109 CFU
basis, with that of commercial pure active dry yeast products. It is also important to know the CFU’s in the products being fed so that
the desired dose is achieved and the expected growth and health responses are observed.
Storage method is very important for maintaining stability and fermentative activity of active dry yeast. When active dry yeast
products are stored in the presence of oxygen or air, their stability is reduced. Therefore, use of vacuum-packaging or storing in
packaging material with inert gas will help ensure optimum stability (Stone, 2006).
Many animal feeds are manufactured and fed as pellets. The pelleting process uses steam (moist heat) to condition the feed prior
to forcing through a pellet die. Moist heat is considered to be one of the most effective methods for killing microorganisms because it
denatures enzyme systems and their metabolic activity within the organism which inhibit their survival. Therefore, when adding
active dry yeast products to animal feeds, there is concern that their viability will be reduced, or potentially eliminated, rendering
them ineffective as probiotics. Very few studies have been conducted to evaluate yeast survival under various feed manufacturing
conditions. Dry, active Sacchromyces cerevisiae (7% moisture) survival was unaffected when heated at 85° C for 8 min, but when the
moisture was increased to 19%, only 0.1% survived (Lewis, 1990). Stone (1998) showed similar results when evaluating pelleting
conditions on Sacchromyces cerevisiae, Kluyveromyces marxianus, and Candida utilis survival. Studies have determined viable yeast cell
counts under different moisture and temperature conditions, and moist heat and dehydration were shown to be the primary factors
causing yeast cell death (Bayrock and Ingledew, 1997a,b). These results, along with those reported by Aguirre-Guzman et al. (2002),
suggest that survival of unprotected yeast cells would be expected to be low after the typical pelleting process. Therefore, mea-
surement of both cell viability and cell vitality must be considered when evaluating the functionality of yeast cells in pelleted feeds,
and the advantages and disadvantages of different methods of evaluating viability and vitality have been compared (Kwolek-Mirek
and Zadrag-Tecza, 2014).
3.2. Yeast culture
Yeast cultures (AAFCO – 96.8 Yeast Culture; IFN 7-05-520 Yeast culture dehydrated) are unique among yeast products because
they contain a combination of yeast biomass and fermentation metabolites produced during specific fermentation processes. While
they contain some residual viable yeast cells, they are not a substantial source of yeast biomass. To produce yeast cultures, a specific
culture media is inoculated with live yeast cells and allowed to ferment under specific conditions, upon which the entire fermented
media is subsequently dried. As yeast cells ferment the sugars present in the culture media, they produce a wide variety of metabolic
products including peptides, alcohols, esters, and organic acids. However, the composition of the various metabolites produced
depends on the composition of the fermentation media used and the fermentation conditions. Therefore, yeast cultures are unique
feed additives that contain several undefined metabolites that may have beneficial nutritional and health effects for animals, but are
not considered to be a nutrient source in animal feeds.
3.3. Nutritional yeast products
Dried yeast (AAFCO – 96.1 Primary Dried Yeast or Dried Yeast; IFN 7-05-533 Yeast primary dehydrated), brewer’s dried yeast
(AAFCO – 96.4 Brewers Dried Yeast; IFN 7-05-527 Yeast brewers dehydrated), brewer’s liquid yeast (AAFCO – 96.10 Brewers Liquid
Yeast; IFN 7-20-878 Yeast brewers liquid), Torula dried yeast (AAFCO – 96.7 Torula Dried Yeast or Candida Dried Yeast; IFN 7-05-
534 Yeast torula dehydrated), and whey yeast are considered to be nutritional yeast products because they consist of yeast biomass of
dead yeast cells and are fed for their nutritional value (Dubey et al., 2010). Dried yeast is expensive and is mainly used in the food
62
industry as a nutritional supplement (Dubey et al., 2010). Brewer’s yeast is commonly used in the animal feed industry as a specialty
amino acid, vitamin and mineral supplement. Torula yeast (Candida utilis) also contains high concentrations of protein, minerals, and
vitamins and it is used as an additive to a wide variety of processed foods (Bekatorou et al., 2006). Historically, Torula yeast was also
used in animal feeds, but today, most of it is used by food manufacturers. Although whey yeast was commonly available and used in
the feed industry for many years, very little is produced and available today.
As shown in Table 1, the crude protein content of Brewer’s yeast is about 46.5% which is similar to that in dehulled soybean meal.
However, about 20% of the crude protein in yeast corresponds to nucleic acids (6–8% of total composition; Reed and Nagodawithana,
1991), which limits the amount of yeast products that can be fed to animals. Furthermore, although the total lysine, methionine, and
threonine content of Brewer’s yeast products is superior to that found in dehulled soybean meal, the standardized ileal digestibility
(swine) of all indispensable amino acids in Brewer’s yeast is substantially less than in dehulled soybean meal. Brewer’s yeast contains
about twice the amount of lipid (ether extract) and half the amount of neutral detergent fiber than dehulled soybean meal, which
contribute to the greater metabolizable energy content (swine) in Brewer’s yeast. Furthermore, Brewer’s yeast contains much greater
phosphorus content, and is a rich source of B vitamins (data not shown; INRA, 2004) compared with dehulled soybean meal.
However, the benefits of high B vitamin content in these yeast products are generally not captured in commercial animal feeds
because of uncertainty of their quantity and bioavailability, and also because diets are routinely supplemented with vitamin-trace
mineral premixes, which are formulated to contain concentrations of these micronutrients substantially greater than the animal’s
requirements.
Most of the nutrients present in whole yeast cells (i.e. brewer’s yeast or active dry yeast) are derived from within the cell.
Therefore, yeast cells must be lysed to release these nutrients for digestion and absorption by the animal. Cell lysis can be accom-
plished by microorganisms in the gastrointestinal tract of animals that secrete proteases and glucanases to hydrolyze the outside of
the yeast cells and release their contents. Alternatively, enzymes within live yeast cells can cause autolysis of the cell wall to release
these nutrients. The differences in digestibility and bioavailability of amino acids and vitamins derived from live versus dead yeast
cells is unknown, but may be greater for live yeast cells because of the higher drying temperatures used to produce dead yeast
products. It has been well documented that use of excessive temperatures when drying feed ingredients can lead to the formation of
Maillard products which reduce amino acid availability (Almeida et al., 2013), and can accelerate lipid peroxidation which can
reduce biological activity of fat soluble vitamins (Shurson et al., 2015).
Technologies are emerging to produce nutritional yeast products from low-value, non-food lignocellulosic biomass for use in
aquaculture feeds (Overland and Skrede, 2016). The general process of using lignocellulosic biomass for yeast production involves
four major steps including pre-treatment, enzymatic hydrolysis, fermentation, and downstream processing (Overland and Skrede,
2016). Studies have shown that some enzymes are capable of cleaving glycosidic bonds in recalcitrant polysaccharides present in
lignocellulosic biomass (Lane and Morrissey, 2010; Horn et al., 2012), and sugars in this biomass can be converted to yeast biomass
by using suitable yeast strains that may be genetically modified (Attfield and Bell, 2006; Jeffries and Jin, 2004; Nielsen et al., 2013).
While nutritional yeast products derived from lignocellulosic biomass are not commercially available, this technology is promising
and would create more high-value, sustainable nutritional yeast products.
Table 1
Comparison of the nutritional composition and standardized ileal digestibilitya for swine of Brewer’s yeast with dehulled soybean meal (INRA,
2004).
Component, % Brewer’s yeast Dehulled soybean meal
Dry matter 93.30 ± 2.5 87.8 ± 0.6

Crude protein 46.5 ± 4.3 45.3 ± 1.0
Ether extract 3.9 ± 1.1 1.9 ± 0.4
Starch 1.0 0.0
Neutral detergent fiber 6.2 ± 8.7 12.2 ± 1.7
Acid detergent fiber 1.8 ± 2.4 7.3 ± 1.9
Ash 7.1 ± 1.1 6.4 ± 0.5
Arginine 2.06 (78) 3.36 (94)
Cysteine 0.29 (49) 0.67 (86)
Histidine 0.98 (77) 1.20 (91)
Isoleucine 1.99 (72) 2.09 (90)
Leucine 2.76 (73) 3.34 (89)
Lysine 2.85 (74) 2.78 (90)
Methionine 0.70 (69) 0.64 (92)
Phenylalanine 1.61 (66) 2.28 (91)
Threonine 2.01 (66) 1.77 (87)
Tryptophan 0.49 (55) 0.59 (89)
Valine 2.20 (66) 2.18 (88)
Calcium 0.32 ± 0.28 0.34 ± 0.09
Phosphorus 1.16 ± 0.16 0.62 ± 0.05
Metabolizable energy (kcal/kg) for swine 3466 3203
a
Values in parentheses indicate standardized ileal digestibility of amino acids for swine (INRA, 2004).
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3.4. Specialty yeast products
Several specialty yeast products are used in various amounts, and for unique nutritional applications in the feed industry in-
cluding irradiated yeast, selenium yeast, chromium yeast, and Phaffia yeast.
3.4.1. Irradiated yeast

Irradiated yeast (AAFCO – 96.3 Irradiated Dried Yeast, Irradiated __ Dried Yeast; IFN 7-05-529 Yeast irradiated dehydrated)
contains ergosterol which can be converted to vitamin D2 (ergocalciferol) when irradiated with ultraviolet light (Hohman et al.,
2011). At one time, irradiated yeast was an important source of vitamin D activity, but less expensive synthetic vitamin D3 (cho-
lecalciferol) has replaced it in the market.
3.4.2. Selenium yeast

The composition, quality, analysis and safety of selenium yeast (AAFCO – 57.163 Selenium Yeast) has been well described
(Schrauzer, 2006). Selenium yeast is commercially produced and marketed as a highly bioavailable form of Se (selenomethionine),
and has unique effects on metabolism, improves health status of animals, and increases Se content in meat and eggs compared to
feeding inorganic forms (Mahan, 1999). Although brewer’s yeast contains 1–2 mg/kg selenium, some commercial high selenium
yeasts may contain up to 2000 mg/kg Se.
3.4.3. Chromium yeast

Chromium yeast contains trivalent chromium complexed with biologically active peptides, amino acids, and niacin, and appears
to function as a “glucose tolerance factor” that acts as a physiological enhancer of insulin activity to improve carbohydrate meta-
bolism (Lukaski, 1999). Limited studies have been conducted to determine the biological mechanism and effects of chromium yeast
on animal growth performance, carcass composition, and reproduction. Mowat et al. (1993) showed that supplementing cattle diets
with chromium yeast after they were subjected to shipping stress, reduced morbidity and improved growth performance. In pigs,
Lindemann et al. (2008) compared the relative bioavailability and safety of four organic forms of chromium, and showed that Cr
bioavailability was greatest for Cr tripicolinate, followed by Cr methionine, Cr yeast, and Cr propionate, and growth rate of pigs fed
Cr yeast was increased compared with those fed the control diet.
3.4.4. Phaffia yeast

Phaffia yeast (Phaffia rhodozyma; AAFCO – 87.36 Phaffia Yeast) produces a red pigment or carotenoid (astaxanthin), and is used
in trout and salmon feeds to enhance the red pigmentation in fillets (European Commission of Health and Consumer Protection
Directorate-General, 2003). Although this source of astaxanthin is more expensive than synthetic forms of this carotenoid, it may
have greater bioavailability because it is associated with an organic matrix (European Commission of Health and Consumer
Protection Directorate-General, 2003).
3.5. Fractionated yeast products
Yeast condensed solubles (AAFCO – 96.9 Molasses Yeast Condensed Solubles; IFN 5-14-009 Sugarcane molasses yeast solubles
condensed), hydrolysates (AAFCO – 96.12 Hydrolyzed Yeast; T96.13 Molasses Hydrolyzed Yeast), extracts (AAFCO – 96.11 Yeast
Extract), and cell walls comprise another category of ingredients produced and used in the food and feed industry. Yeast autolysates
consist of lysed whole yeast cells (using acids or enzymes or high salt solution to rupture cells caused by increased osmotic pressure),
and contain both intracellular and cell wall fractions. Yeast extracts consist of only the intracellular components, while the yeast cell
wall products consist only of the carbohydrates found in the cell wall. Yeast autolysates and extracts are used primarily by the food
industry for flavor enhancement of foods (Hassan, 2011). Yeast extracts are also used as microbial stimulants in fermentation and
pharmaceutical industries, as well as by microbiologists to optimize bacterial growth and in laboratory growth media (Hassan, 2011).
The carbohydrate content and composition of yeast varies depending on the growing conditions, and are often classified as either
intracellular or cell wall carbohydrates (Halasz and Lasztity, 1991). Intracellular carbohydrates are primarily comprised of glycogen
and trehalose, which serve as the major energy reserves for yeast cells. Baker’s yeast has been reported to contain 16–20% glycogen
and 6–10% trehalose (Sols et al., 1971). The cell wall represents about 15–20% of the dry weight of yeast cells, and the main
polysaccharide fractions are β-glucans and mannans.
Glucans are the primary polysaccharide component in Saccharomyces cerevisiae, and are a highly insoluble part of the support
structure of the cell wall. The inner layer of yeast cell walls consists of insoluble β-glucan (30–35%), the middle layer is soluble β-
glucan (20–22%), and the external layer consists of glycoprotein (30%; Tokunaka et al., 2002). The structure of glucans consist of β-
1,3 glucans with β-1,6 branch linkages, and both endo and exo β-1,3 glucanases are produced during yeast autolysis (Halasz and
Lasztity, 1991). β-glucans are becoming very popular in the animal feed industry because of their potential beneficial effects on
animal health and growth performance. They have been shown to adsorb or bind toxins, viruses, and pathogenic bacteria (Vetvicka
et al., 2014). Immune cells (macrophages) have receptors for β 1,3/1,6 branched glucans. Their mode of action is well documented in
human nutrition and medicine (Rop et al., 2009), and there is evidence that dietary β-glucans may improve immune-competence in
young animals (Saeed et al., 2014), as well as in fish and shrimp.
Mannans are considered to be the second most important component of yeast cell walls, and are linked by α-1,6 bonds in a branch
structure, with side chains consisting of mannose units linked to the backbone with α-1,2 bonds (Halasz and Lasztity, 1991).
64
Phosphorus is connected to mannans, and the amount varies from 0.04 to 4.4%. Mannanoligosaccharides (MOS) serve as prebiotics,
or sources of nutrients for select microbes in the gastrointestinal tract, that can lead to providing a probiotic effect (Spring et al.,
2015). Spring et al. (2015) reviewed results from 733 published trials that evaluated the effects of feeding MOS to companion
animals, horses, rabbits, poultry, pigs, calves, and various aquaculture species, and showed generally positive improvements in body
weight gain and feed conversion, while reducing mortality. These beneficial responses are a result of MOS binding and limiting the
colonization of pathogens in the gastrointestinal tract, improving the integrity of the intestinal mucosa, modulating immune system
activity, and may be involved in antioxidant and anti-mutagenic defenses (Spring et al., 2015).
Chitin is another polysaccharide found in yeast cell walls, but its concentration is much less than for glucans and mannans (Halasz
and Lasztity, 1991). Chitin is a polymer of glucosamine found in shells or walls of invertebrates, fungi and yeast, and is the main
component of crustacean exoskeletons. However, its effects on animal nutrition and health are not well studied.
Yeast is also a concentrated source of nucleotides and proteins. Hydrolytic enzymes (nucleases and proteases) are located in the
general matrix of the yeast cell to degrade these nucleotides and proteins by autolysis. Nucleases degrade nucleic acids, DNA, and
RNA into nucleotides, and proteases degrade proteins into peptides and amino acid derivatives (Reed and Nagodawithans, 1991;
Sommer, 1998). Concentrations of total nucleic acids have been reported to be between 3.3–9.5% (Bacha et al., 2013) and 7–12%
(Halasz and Lasztity, 1991) in yeast. Knowing the nucleic acid content of yeast and yeast containing products is important because
there is increasing evidence that the supplementation of nucleotides in diets of monogastric animals may affect intestinal morphology
and function, immune response, composition of intestinal microbiota, liver function and morphology, as well as growth performance
(Sauer et al., 2011).
3.6. Yeast and yeast components in corn ethanol co-products
Large quantities of various co-products are produced after grain (e.g. maize) is fermented with yeast in ethanol production. These
co-products include distillers dried solubles (AAFCO 27.4; IFN 5-02-844 Maize distillers solubles dehydrated), distillers dried grains
(AAFCO – 27.5; IFN 5-02-842 Maize distillers grains dehydrated), distillers dried grains with solubles (AAFCO – 27.6; IFN 5-02-843
Maize distillers grains with solubles dehydrated), condensed distillers solubles (AAFCO – 27.7; IFN 5-12-211 Maize distillers solubles
condensed), distillers wet grains (AAFCO – 27.8; IFN 5-16-149 Maize distillers solubles condensed), deoiled maize distillers dried
grains with solubles (AAFCO – T27.9), yeast for production of distiller’s products (AAFCO – 73.100), and grain distillers dried yeast
(AAFCO – 96.5). All of these co-products are produced in various quantities by the beverage and fuel ethanol industries and fed to
food producing animals (Shurson, 2017). These co-products are produced using different processes involving different strains of
Saccharomyces cerevisiae yeast, which is present in various quantities in these co-products, contributes to their nutritional compo-
sition, and may provide functional properties and health benefits when fed to animals.
The composition and functional properties of yeast in these co-products is not well established because of a lack of standardized
analytical methods to determine the dead yeast and its chemical components in these ingredients. Published estimates of the con-
tributions of yeast to DDGS biomass and crude protein are extremely variable. In 1944, Bauernfiend et al. (1944) proposed that dead
yeast cell content could be estimated by using a hemocytometer to count yeast cells in thin stillage, condensed distillers solubles, or
dried solubles, and determined that distillers dried solubles contained about 4 × 109 cells per gram. This estimate was then compared
with the cell count in dried yeast to calculate that about 20% of the weight of dried solubles is contributed from yeast. However, this
method is not applicable for estimating the yeast content in DDGS because it relies on separating or assuming the proportion of
solubles that is added to the grains fraction prior to manufacturing DDGS. Ethanol plants vary in the proportion of condensed solubles
blended with the coarse grains fraction prior to manufacturing DDGS as defined by the AAFCO definition 27.6.
Ingledew (1999) used a mass balance approach and numerous assumptions involving ethanol production processes in the late
1990’s, to estimate the amount of yeast that was contributed to the biomass and crude protein content of DDGS. Based on the large
number of assumptions used in these calculations, and the fact that the ethanol and co-product production processes have changed to
improve the efficiency of production since the date of this publication, estimates of the proportion yeast in DDGS mass from this study
are unreliable.
Belyea et al. (2004) used a different, and much simpler, approach to estimate the yeast contribution to total protein in DDGS.
These researchers calculated the average ratio of amino acid concentrations (dry matter basis) of yeast and DDGS and suggested that
about 50% of DDGS protein was derived from yeast. However, this method has several serious flaws. First, there is no consideration
given for the crude protein contribution from corn in the total crude protein content of DDGS. Second, nonessential amino acid
content was not considered in the calculations. Finally, the average ratio of amino acids were based on the amino acid ratios in DDGS,
not yeast.
Most recently, Han and Liu (2010) developed a multiple linear regression model that includes the relative percentage changes in
the amino acid profile throughout the entire DDGS production process to determine the contribution of yeast protein to total DDGS
protein content:
Y = AX1 + BX2 + C
where Y = the relative percentage of an amino acid in an ethanol production process stream, X1 = the relative percentage of an
amino acid in ground corn, X2 = the relative percentage of an amino acid in yeast, A = a constant indicating the amount of amino
acid contribution from corn, B = a constant indicating the amount of amino acid contribution from yeast, and C = a constant for the
intercept on the Y axis. From their analysis, the A, B, and C variables varied substantially among various production streams, but only
65
slightly among sample sources. Based on this equation, which accounts for the amino acid composition of corn and yeast as in-
dependent variables, they estimated that yeast accounts for 20% of the crude protein content in DDGS and corn contributed 80% of
the total protein.
While none of these methods directly measure the yeast content in DDGS, the calculation of Han and Liu (2010) likely provides
the most accurate estimate of all methods developed to date. Unlike methods used by Bauernfiend et al. (1944) and Belyea et al.
(2004), the multiple linear regression model accounts for the amino acid profiles in both corn and yeast, includes all amino acids, is
based on relative percentages of amino acids rather than absolute concentrations of amino acids, and can be used to estimate yeast
content in intermediate process streams as well as in DDGS. However, amino acid profiles vary among corn varieties and crop year,
and can also vary among yeast strains. Furthermore, the new production processes being implemented in several U.S. ethanol plants
will likely change the amino acid concentrations in various process streams which may lead to questionable estimates. Finally, while
it may be appropriate to estimate yeast content in DDGS on a protein or amino acid basis, the greater potential nutrition and health
benefits of yeast in corn co-products may be their β-glucan, mannanoligsaccharide, and nucleotide content, which cannot be esti-
mated using this method.
Like estimates of yeast content in DDGS, there are wide discrepancies among published studies on the β-glucan concentrations in
DDG. Lim et al. (2009) estimated that DDGS contained 0.57% β-glucans, whereas Kim et al. (2008) reported that the average total
glucan content in DDGS was 21.2%. It is likely that this wide difference is due to differences in analytical procedures used and the
actual composition of the β-glucans measured. Alizadeh et al. (2016) reported that the concentration of mannose and nucleotides in
wheat-corn DDGS was 1.6% and 0.13%, respectively. As a result, there is a lack of standardized analytical methods and the inherent
difficulty of determining yeast content and derivatives in fermented grain co-products.
4. Animal responses to diets containing live yeast, yeast cell walls, and DDGS
Except for various methods for determining live yeast concentrations and viability, there are no methods available for accurate
Table 2
Summary of positive effects of dietary yeast supplementation on animal health, nutrition and growth performance (adapted from Vohra et al., 2016).
Animal species Yeast probiotic responses References
Ruminants Improves growth of lactic acid utilizing microorganisms and reduces Chaucheyras-Durand et al. (1996), Mathieu et al. (1996), Marden
lactic acid production or accumulation et al. (2008), Robinson (2010)
Balances rumen pH by modifying lactic acid and volatile fatty acid Lila et al. (2004), Desnoyers et al. (2009), Hucko et al. (2009),
concentrations in the rumen to reduce the risk of rumen acidosis and Thrune et al. (2009)
related digestive disorders
Stimulates growth of cellulolytic and fibrolytic microorganisms and Wohlt et al. (1988), Callaway and Martin (1997), Miller-Webster
increases fiber intake and digestion et al. (2002), Mosoni et al. (2007), Di Francia et al. (2008),
Chaucheyras-Durand et al. (2008), Guedes et al. (2008), Marden
et al. (2008)
Removes oxygen and increases total rumen microflora Jouany (2006), Chaucheyras-Durand and Durand (2010)
Improves growth performance and milk production Lesmeister et al. (2004), Haddad and Goussous (2005), Stella et al.
(2007), Masek et al. (2008), Moallem et al. (2009), Desnoyers et al.
(2009), Pal et al. (2010), Bitencourt et al. (2011)
Reduces methane emissions Lynch and Martin (2002), Lila et al. (2004), Mwenya et al. (2004)
Horses Reduces lactic acidosis, colitis, laminitis, and enterotoxemia Medina et al. (2002), Bertin and McCartney (2005), Desrochers
et al. (2005)
Improves nutrient digestibility Glade (1991), Morgan (2007), Bertin and McCartney (2005),
Jouany et al. (2008), Agazzi et al. (2011)
Enhances enzyme production in the lower gastrointestinal tract Bertin and McCartney (2005), Chaucheyras-Durand and Durand
(2010)
Pigs Improves microflora balance in the gastrointestinal tract Chaucheyras-Durand and Durand (2010)
Participates in maturation of tissues in the gastrointestinal tract Shen et al. (2009)
Modulates immune response to reduce enteric pathogens Shen et al. (2009)
Reduces post-weaning diarrhea Chaucheyras-Durand and Durand (2010)
Improves growth rate and feed conversion Shen et al. (2009)
Poultry Modulates intestinal micoflora Hassanein and Soliman (2010), Javadi et al. (2012)
Inhibits growth of pathogenic bacteria Line et al. (1998), Hassanein and Soliman (2010)
Improves egg production, egg shell weight and internal egg quality Hassanein and Soliman (2010)
Improves feed intake and feed conversion Shareef and Al-Dabbagh (2009), Hassanein and Soliman (2010)
Reduces plasma cholesterol Hassanein and Soliman (2010)
Improves blood chemistry Shareef and Al-Dabbagh (2009)
Improves carcass characteristics and meat quality of broilers Pelicano et al. (2003)
Aquaculture Improves growth rate of juvenile and adult fish Noh et al. (1994), Tovar-Ramírez et al. (2004)
Improves survival rate, maturation and resistance of Artemia as a fish Patra and Mohamed (2003), Shoja et al. (2012)
food source, against Vibrio infection
Stimulates enzymatic antioxidant responses in farmed fish Tovar-Ramírez et al. (2010)
Stimulates digestive enzymes in seabass larvae Tovar-Ramírez et al. (2010)
Secretes polyamines and enhances maturation of the gastrointestinal Andlid et al. (1998), rez et al. (2004, 2010);
tract
66
quantification of dead yeast and yeast derivatives in animal feeds. It is imperative that methods be developed to quantify yeast
components because of the widespread use of these products as growth and health enhancers in animal feeds. Numerous published
studies have shown that the presence or supplementation of these yeast products in diets for all animal species may provide sig-
nificant benefits. Yeast and yeast-based products are generally fed to animals as live yeast (direct fed microbial; DFM; probiotic),
yeast cell wall components (mannan-oligosaccharides, β-glucans, and nucleotides), ethanol co-products (DDGS, specialty high pro-
tein distillers grains), or a combination of these.
4.1. Probiotic effects of live yeast
The benefits of adding yeasts as probiotics to animal feeds have been reviewed by Vohra et al. (2016; Table 2). The majority of
studies evaluating live yeast as a probiotic have been conducted in ruminants, with fewer studies in horses, swine, poultry, and
aquaculture. However, studies have generally shown that the addition of live yeast may improve fiber digestibility, prevent pathogen
growth, produce antibacterial compounds, and modulate the immune system. Song et al. (2014) reviewed several studies that
evaluated the immune responses of finfish and shellfish fed probiotics, and indicated that these compounds directly activate the
innate immune system by interacting with pattern recognition receptors, as well as being involved in microbe associated molecular
patterns to activate innate immune cells. Other reviews showing the beneficial effects of supplementing aquaculture feeds with
prebiotics and probiotics have been published (Akhter et al., 2015; Hai, 2015; Song et al., 2014). However, in aquaculture, the
beneficial effects of probiotics fed to tilapia are not always observed, but responses appear to be better understood than for other fish
species (Hai, 2015). Furthermore, Broadway et al. (2015) reviewed the literature on feeding live yeast and yeast cell wall supple-
ments to food-producing animals and noted that yeast cell wall components interact directly with immune cells, bind bacteria to
prevent colonization of pathogens, may possess antioxidant and antitumor properties, enhance growth performance, and alter me-
tabolism.
4.2. Health benefits of yeast cell wall components
4.2.1. Mannanoligosaccharides
Yeast cell walls are concentrated sources of mannan-oligosaccharides (MOS), and several commercial feed additive products
containing concentrated amounts of MOS have been fed to animals for many years. Spring et al. (2015) reviewed results from 733
published trials that evaluated the effects of feeding MOS to companion animals, horses, rabbits, poultry, pigs, calves, and various
aquaculture species, and showed generally positive improvements in body weight gain and feed conversion, while reducing mor-
tality. Meta-analysis studies conducted by Hooge (2004a,b) showed that the magnitude of growth and feed conversion improvements
in broilers (Table 3) and turkeys (Table 4) are relatively small and inconsistent, but more than half of the studies reported substantial
reductions (average of 16–25%) in mortality. Similarly, Miguel et al. (2004) summarized 54 treatment comparisons obtained from
published studies to calculate the relative percentage improvement in growth performance in nursery pigs fed diets containing MOS
compared with those fed control diets (Table 5). The average percentage of improvement growth rate and feed conversion from
feeding MOS to nursery pigs was greater than the magnitude of improvement reported by Hooge (2004a,b) for broilers and turkeys,
but responses were again inconsistent among studies. In pigs, improvements in growth performance from supplementing diets with
MOS supplementation were only observed in low hygiene environments and not in high health swine herds (Halas and Nochta, 2012).
In fish, while MOS supplementation has been shown to improve fish growth performance in several studies (Table 6), the majority of
studies have reported no change (Torrecillas et al., 2014). Several experiments have evaluated fish survival and disease resistance
(Table 6) from feeding MOS supplemented diets compared with unsupplemented control diets and fish health was improved in the
majority of these studies. These beneficial responses have been attributed to MOS binding and limiting the colonization of pathogens
in the gastrointestinal tract, improving the integrity of the intestinal mucosa, enhancing immune system activity, and may be in-
volved in antioxidant and antimutagenic defenses (Spring et al., 2015). However, these effects are dependent on the molecular
structure of MOS fed, dose and duration of feeding, species, stage of growth, and culture conditions (Torrecillas et al., 2014; Song
et al., 2014).
4.2.2. β-Glucans
In addition to MOS, yeast cell walls contain β-glucans, which are also often classified as prebiotics (Song et al., 2014). β-glucans
are glucose polymers that are present in cell walls of yeast, fungi, and some bacteria, as well as in endosperm cell walls of cereal
Table 3
Meta-analysis summary of growth performance and mortality responses of broilers fed diets containing mannan oligosaccharides compared with feeding antibiotic-free
negative control diets (Hooge, 2004a).
Parameter Average Responsea, % No. Studies Reporting an Increase No. Studies Reporting a Decrease No. Studies Reporting No Change
Body weight gain +1.75 (44) 35 6 3

Feed:Gain −1.89 (43) 5 35 3
Mortality −16.4 (29) 10 18 1
a
Numbers in parentheses indicate number of comparisons.
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Table 4
Meta-analysis summary of growth performance and mortality responses of turkeys fed diets containing mannan oligosaccharides compared with feeding antibiotic-free
negative control diets (Hooge, 2004b).
Body weight gain +2.25 (27) 19 8 0

Feed:Gain −1.55 (24) 11 13 0
Mortality −24.6 (16) 3 9 4
a
Number in parentheses indicate number of trials in the comparisons.
Table 5
Summary of growth performance responses of nursery pigs fed diets containing mannan oligosaccharides compared with feeding control diets (Miguel et al., 2004).
ADG2 +4.12 (54) 36 17 1

ADFI3 +2.11 (54) 36 16 2
Feed:Gain −2.29 (54) 12 38 4
a
Numbers in parentheses indicate number of comparisons.
Table 6
Summary of growth performance, survival, and disease resistance responses of fish fed diets containing mannan oligosaccharides compared with feeding
control diets (Torrecillas et al., 2014).
Parameter No. Studies Reporting an Improvement No. Studies Reporting No Change
Growth rate 12 19
Feed:Gain 6 18
Survival 9 3
Disease resistance 6 2
grains such as oats and barley (Volman et al., 2008). However, the structure of β-glucans varies among sources, and may affect their
physiological functions. The β-glucans in yeast cell walls primarily consists of 1,3 β linkages of glycopyranosyl residues with limited
1,6 β-linked branches, while endosperm cell walls of oats and barley contain unbranched 1,3 and 1,4 β-linked glycopyranosyl
residues (Estrada et al., 1997; Brown and Gordon, 2003; Brown et al., 2003). In addition, β-glucans vary among sources in their
solubility, molecular weight, tertiary structure, extent of branching, polymer charge, and conformation (Volman et al., 2008).
Several studies have shown that β-glucans enhance the immune response in leukocytes and epithelial cells and improve survival
of the host after a pathogen infection (Volman et al., 2008). Dietary β-glucans modulate the intestinal mucosal immune response
through direct effects on the Peyer’s patches and intestinal intraepithelial lymphocytes (Suzuki et al., 1990; Tsukada et al., 2003).
Suzuki et al. (1990) suggested that feeding 1,3 β-glucans improves immune response through receptor-mediated interactions with M-
cells, which are specialized epithelial cells used for transporting macromolecules in the Peyers patches, result in increased cytokine
production and resistance to infection. Orally administered whole glucan particles from yeast may be bound to intestinal macro-
phages and transported to lymph nodes, spleen, and bone marrow where they may be degraded and enter the circulatory system in
low amounts to provide systemic effects (Volman et al., 2008). Regardless of the extent to which yeast β-glucans are absorbed, their
effects on the immune system are deptin-1 dependent, and the deptin-1 receptor is highly expressed in immune cells (i.e. dendritic
Table 7
Summary of studies evaluating immune system effects of supplementing pig diets with β-glucans
(adapted from Vetvicka et al., 2014).
Immune response Reference
No effects Hiss and Sauerwein (2003)

Minimal effect on antibody production Amornlertviman et al. (2008)
Improved hematology Amornlertviman et al. (2008)
Downregulation of IL-6 and TNF-α Li et al. (2006)
Upregulation of IL-10 Clarke et al. (1998)
Decrease in Th-related cytokines Ryan et al. (2012)
Protection against infection Stuyven et al. (2009)
Stimulation of phagocytosis Benova et al. (1992)
Protection against ascariosis Benova et al. (1992)
Increased IgA production Sauerwein et al. (2007)
Stimulation of IL-2 and phagocytosis Vetvicka and Oliverira (2014)
Suppression of TNF-α Vetvicka and Oliverira (2014)
68
cells, neutrophils, eosinophils, macrophages, monocytes, and some T-cells; Volman et al., 2008). Once β-glucan binds to the dectin-1
receptor, intracellular signaling occurs leading to NF-κB activation, which further initiates cytokine production, phagocytosis, and
respiratory burst (Volman et al., 2008).
Vetvicka et al. (2014) reviewed the limited published studies evaluating the effects of dietary β-glucans on pig growth and
immune responses (Table 7). It is assumed that feeding supplemental β-glucans affect the composition of the microbiota in the
gastrointestinal tract, but little information has been published on these effects. Pigs studies evaluating the supplementation of β-
glucans from different sources have shown improvements in growth rate (Dritz et al., 1995; Kogan and Kocher, 2007), reduced peak
glucose flux (Hooda et al., 2010), decreased TH-related ctoking production (Ryan et al., 2012), and protection from an en-
terotoxigenic Escherichia coli infection (Stuyven et al., 2009). Li et al. (2006) suggested that dietary β-glucan supplementation may
suppress cellular immunity which may lead to improved growth performance. Vetvicka et al. (2014) concluded that supplementing
diets with β-glucans may improve pig health and growth performance, but due to the variation in structure of glucan molecules,
molecular weight, and quantities of various contaminating substances, more studies are needed. This conclusion is consistent with
results from another study that evaluated dietary β-1,3/1,6 glucans on growth performance, lymphocyte proliferation, plasma
haptoglobin concentrations, and specific immune responses of pigs, and showed no improvement in growth rate or immune re-
sponses, but a slight improvement in feed intake (Hiss and Sauerwein, 2003).
Ringo et al. (2012) reviewed and summarized 14 studies involving feeding yeast β-glucans to various fish species and reported
improvements in pathogen resistance, growth performance, and survival. Limited studies have been conducted to evaluate the po-
tential growth and immune system benefits of β-glucans in ruminants. Feeding β-glucans from oats improved immune responses in
immunosuppressed cattle, but had no effects in health animals (Estrada et al., 1999). Eicher et al. (2010) fed β-glucans derived from a
yeast-like fungus and suggested that it may alter expression of serum cytokines and alter microflora of the intestinal tract. Uchiyama
et al. (2012) showed that feeding different types and purity of β-glucans increased feed intake, but provided different immune
responses.
4.2.3. Nucleotides
Yeast is also a concentrated source of nucleotides (Bacha et al., 2013). Nucleotides are a class of molecules that are linked together
to form DNA and RNA, and are composed of a phosphate group; adenine, cytosine, guanine, and thymine bases; and a pentose sugar.
In RNA, the thymine base is replaced by a uracil base. Nucleotides have been shown to improve intestinal morphology and function,
immune response, composition of intestinal microbiota, liver function and morphology, as well as growth performance (Sauer et al.,
2011). Studies have shown that nucleotide supplementation in animal diets may be necessary under certain conditions (rapid growth,
reproduction, environmental changes, and disease challenges; Yu, 1998; Carver, 1999). Although limited studies have been con-
ducted, results have generally shown that dietary supplementation with nucleotides improves growth performance of broilers (Rutz
et al., 2006; Esteve-Garcia et al., 2007), gut morphology in rats (Uauy et al., 1990) and pigs (Domeneghini et al., 2004; Martinez-Puig
et al., 2007), and growth performance during an E. coli challenge in pigs (Maribo, 2003). However, other studies have shown that
when animals were not reared under stressful conditions, there was no beneficial effects on growth performance of chickens (Deng
et al., 2005; Jung and Batal, 2012) and pigs (Andres-Elias et al., 2007; Martinez-Puig et al., 2007).
Because most studies have evaluated commercially available yeast products, which contain variable amounts of nucleotides,
viable cells, cell wall components, and media components and metabolites, it is difficult to attribute these positive responses solely to
nucleotides (Sauer et al., 2011). Ringo et al. (2012) reviewed and summarized 15 studies involving feeding various sources of
nucleotides to several fish species and reported consistent improvements in immune responses, pathogen resistance, growth per-
formance, and survival. In contrast, Sauer et al. (2011) summarized 16 published studies of growth responses from feeding various
concentrations of Saccharomyces cerevisiae, yeast culture, and commercial products containing nucleotides to pigs (Table 8), which
provided some evidence in growth performance improvements, but most of the responses showed no effect. It is important to note
that only one of these sources (yeast RNA) contained pure nucleotides, whereas the other products were mixtures of nucleotides with
other compounds. Increased growth performance responses were observed when live Saccharomyces cerevisiae cells, yeast culture, and
some commercial products were fed, but these are not pure sources of nucleotides.
4.2.4. Biopeptides
Historically, the cost of adding yeast extracts as a source of biopeptides to animal feeds was prohibitive. However, increased
Table 8
Summary of the number of published studies showing no change, increase, and decrease in growth performance responses of pigs fed various concentrations and
sources of yeast-derived nucleotides (adapted from Sauer et al., 2011).
Nucleotide source Average Daily Gain Average daily Feed Intake Gain:Feed
No change Increase Decrease No change Increase Decrease No change Increase Decrease
S. cerevisiae culture 2 0 1 2 0 1 2 0 1
Live S. cerevisiae cells 0 1 0 0 1 0 0 0 0
Yeast culture 1 2 0 1 2 0 1 2 0
Yeast RNA 1 0 0 1 0 0 1 0 0
Commercial products 4 2 0 4 1 0 4 1 0
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demand for yeast cell wall-based products has increased the availability and decreased costs of yeast extracts. Limited studies have
been conducted to determine the potential health benefits of biopeptides on animal growth performance and health, but one study
has shown an improvement in feed conversion when supplementing broiler starter diets with yeast-derived biopeptides (Dawson,
2001).
4.2.5. Mycotoxin binding

Yeast and yeast cell wall derivatives appear to have some ability to bind mycotoxins (aflatoxins, ochratoxin A, T-2 toxin, and
zearalenone) and minimize their adverse effects on animal health and performance (Shetty and Jespersen, 2006). Some studies have
shown that removal of mycotoxins is by adhesion to cell wall components (i.e. mannans and β-glucans), and not by covalent binding
or metabolism, because dead yeast cells do not lose their binding ability (Baptista et al., 2004; Celyk et al., 2003; Raju and
Devegowda, 2000; Santin et al., 2003).
4.2.6. Immune responses from feeding DDGS

As previously discussed, yeast and yeast components are present in DDGS and other co-products produced by the ethanol in-
dustry, but there is limited information and lack of reliable analytical methods to determine the concentrations of biologically active
yeast-derived compounds in these feed ingredients. However, limited studies have shown some evidence of positive animal health
responses from feeding DDGS diets, but these responses have been inconsistent. Presumably, these inconsistent responses are a result
of dietary DDGS inclusion rate, concentrations of bioactive yeast components, animal species, and type of disease challenge.
In swine, the addition of 10% DDGS to the diet of growing pig experimentally challenged with Lawsonia intracellularis reduced the
prevalence, length, and severity of intestinal lesions compared with infected pigs fed the corn-soybean meal control diet (Whitney
et al., 2006). Recovery time was increased in young pigs challenged with pathogenic Escherichia coli and fed a DDGS diet (Perez,
2010).
However, in poultry, feeding a DDGS diet did not prevent or reduce an Eimeria acervulina (a prevalent species of coccidia)
infection, but changes in cecal microbiota were observed suggesting that feeding DDGS may be beneficial for intestinal health (Perez
et al., 2011). Necrotic enteritis (caused by Clostridium perfringens types A and C), reduces feed intake and nutrient absorption, causes
diarrhea, and severe necrosis of the intestinal tract of poultry. When broilers were infected with Clostridium perfringens, feeding DDGS
diets has either increased the severity of intestinal lesions (Barekatain et al., 2013), or had no effect (Perez et al., 2011). It is likely
that other dietary components, such as protein source and digestibility and the use of enzymes play a role in the susceptibility and
severity of necrotic enteritis infections in poultry. Alizadeh et al. (2016) fed various yeast-derived products and 10% wheat-corn
DDGS to broilers and observed down-regulation of receptors and cytokines in spleen and ceca of birds fed diets containing yeast-
derived products and suggested that these products do not have immune stimulatory effects in healthy birds.
Lim et al. (2007) fed Nile tilapia diets containing up to 20% DDGS to Nile tilapia (Oreochromis niloticus) infected with Streptococcus
iniae and observed no changes in growth performance, body composition, hematology, immune response, and resistance to infection.
However, in a subsequent study, feeding DDGS diets to channel catfish (Ictalurus punctatus) improved resistance to an Edwardsiella
ictaluri challenge (Lim et al., 2009).
5. Methods to measure yeast and yeast derivative concentrations
5.1. Direct measurement of yeast cells
In a review article published in 1996, Hobson et al. (1996) indicated that there is widespread need for commercial in-
strumentation that can rapidly and inexpensively detect microbial contamination in food, industrial wastewater, and clinical samples.
This continues to be a relevant statement today regarding the need for commercially available instrumentation for the determination
of yeast and yeast derivatives in feed additives and ingredients.
The only methods that have been developed to quantify yeast composition in feed additives and feed ingredients apply only to
viable yeast products (active dry yeast). The AOAC (1997) has an official method (977.02) for determination of yeast and mold
counts in foods and feeds, but does not measure yeast cell remnants. Several methods have been developed and evaluated to de-
termine concentration and viability of pure yeast samples including light and fluorescence microscopy using a hemocytometer, flow
cytometry, and fluorescence microscopy (Smart, 2003). Hemocytometry is relatively inexpensive and simple to use, but it is tedious
and prone to human error which leads to high variability in estimates (Szabo et al., 2004). Flow cytometry provides automated data
acquisition, but has the disadvantages of being expensive and requiring trained technicians to properly operate and clean the
equipment to avoid cross-contamination between samples (Michelson, 1996). Fluorescence microscopy quantification of yeast has
been evaluated in several studies involving pure yeast samples (King et al., 1981; McCaig, 1990; Slater, 1976; Zanducke et al., 2003).
Manual counting methods using a hemocytometer to determine yeast content in corn mash provide inconsistent results because
debris in corn mash can interfere with flow cytometry systems and cause nonspecific fluorescent signals (Taylor et al., 2001). More
recently, Chan et al. (2011) developed and evaluated a simple imaging cytometry method to determine the concentration and
viability of fermenting yeast in corn mash from bioethanol production. This method uses an automated cell counter, dilution buffer,
and staining solution to accurately detect viable and non-viable yeast content by fluorescence, but has not been evaluated for use in
dried corn co-products.
Manual yeast plate counts appear to be the most accurate method because it determines the actual amount of live, viable yeast
cells in a product. In brief, this method involves serial dilution of a suspension of the yeast-containing product in water, followed by
70
plating the different dilutions on nutrient media for about 3 days, to allow them to grow and multiply. The number of yeast colonies
are counted on each plate and reported as CFU’s per gram.
Methylene blue staining has been the standard method for staining nonviable yeast for manual counting, but it provides in-
consistent concentration and viability results because it is difficult to identify and distinguish nonviable yeast from debris in corn
mash (Abbott and Ingledew, 2004; Boyd et al., 2003; Deere et al., 1998; Trevors et al., 1983; Vairo, 1962). This method requires
destruction of the yeast cell wall in order for the stain to enter the cell. If yeast cells die from denaturation of enzyme systems due to
heat, and the cell wall is unaffected, the dead cell cannot absorb the stain and the cells will appear to be viable when they are not.
Methylene blue staining for counting yeast cells is also time consuming, accuracy varies among technicians, and is difficult to
automate (Boyd et al., 2003; Deere et al., 1998; Mills, 1941; Taylor et al., 2001).
An alternative approach is to determine the metabolic activity of yeast by directly measuring the end-products from metabolism
or disappearance of substrate. Unfortunately, this approach is not as reliable as the yeast plate count method because there can be
substantial interference with metabolites produced by other microorganisms, such as bacteria. Procedures may involve using a simple
manometric apparatus to determine carbon dioxide production, or more complex methods such as monitoring glucose disappearance
or ethanol production.
5.2. Methods to determine yeast viability and vitality
Yeast viability can be described as the percentage of live yeast cells relative to total yeast cells, whereas cell vitality refers to the
physiological capabilities of yeast cells (Kwolek-Mirek and Zadrag-Tecza, 2014). Several active dried yeast products are commercially
produced and marketed for use in animal feeds, and the number of colony forming units of these products determines their viability,
efficacy, and shelf-life (Poppy, 2008). Yeast cells are often included in commercial probiotic animal feed products, and their con-
centration and vitality are important to determine their potential effectiveness. Therefore, it may be desirable to not only determine
the concentration of yeast cells in co-products and yeast-containing feed additives, but also their vitality. Kwolek-Mirek and Zadrag-
Tecza (2014) compared the advantages and disadvantages of using different methods to determine viability and vitality of yeast cells.
They showed that the results varied among methods used and that no single method is adequate to quantify both viability and
vitality. When storing baker’s yeast for up to 16 days, cell death rate increases markedly, and using a fluorochrome technique that
includes primuline, acridine orange or acriflavine as fluorochromes, was shown to be the most effective method for determining the
proportion of dead cells (Parkkinen et al., 1976). Bovill et al. (2001) compared the recovery and effectiveness of probiotic yeast
species differentiation using six different selective media. These researchers indicated that isolation and enumeration of yeasts from
samples containing molds is a significant problem because their presence quickly obscures the surface of agar plates. As a result, this
problem would pose a significant challenge to using these methods to quantify yeast in ethanol corn co-products because of the likely
presence of molds in co-products. Poppy (2008) conducted a study to compare the use of 8 different methodologies to determine yeast
viability in active dry yeast, mixtures of dried viable yeast diluted with a cereal carrier, mineral blends with active dry yeast,
complete feed of grain and forage, and yeast culture. He concluded that there were no differences among methods, except for using
the Bacteriological Analytical Manual DG18 method from FDA, and the methods can be used interchangeably. However, Slaughter
and Nomura (1992) evaluated the potential use of intracellular glycogen and trehalose concentrations as predictors of yeast viability
and showed that while there is a correlation between yeast viability and acid soluble glycogen or trehalose of yeast cells, this
association is specific to experimental conditions and storage carbohydrate content of cells and cannot be used to predict yeast
viability without prior knowledge of storage conditions of yeast.
5.3. Methods to determine yeast and yeast-derivatives in DDGS
Bauernfiend et al. (1944) defined corn DDGS as a grain-yeast concentrate. Although a portion of the total DDGS protein is derived
from yeast, the estimates obtained and the methods used are highly variable and insufficient. As a result, there are no well-defined
analytical methods to determine the dead yeast and its chemical components in distillers co-products. Ingledew (1999) estimated that
3.9% of the total DDGS biomass was yeast, and contributed 5.3% of the total protein as yeast protein. Additional attempts to estimate
yeast content in distillers dried grains with solubles (DDGS) have been made by calculating the proportional contributions of amino
acids from yeast and corn to total amino acid content in DDGS (Belyea et al., 2004; Han and Liu, 2010). In the study by Belyea et al.
(2004), the ratios of essential amino acid concentrations in DDGS and yeast varied among amino acids (ranging from 0.45 to 0.70
with an average of 0.55), but suggested that about half of DDGS protein is derived from yeast. However, this approach ignored the
effect of amino acid composition in corn, and as a result, it is not an accurate assessment of yeast content. Han and Liu (2010) used a
more detailed approach to estimate yeast content in DDGS by conducting a multiple linear regression analysis using the relative
percentage of individual amino acids in corn and yeast, relative to total amino acid of DDGS. From their analysis, they suggested that
about 20% of the protein in DDGS is derived from yeast, with the remaining 80% derived from corn. However, the authors ac-
knowledged that this estimate does not infer that yeast biomass contributed to 20% of DDGS biomass. Therefore, this method is not
an accurate approach for estimating yeast content in fermented corn co-products.
Unique chemical characteristics of yeast cell walls, such as mannan content, can be used as a proxy to more accurately estimate
the yeast content in maize co-products. Recent unpublished data (K. Englyst, Englyst Carbohydrates, Ltd., Southampton, UK) showed
that the average mannan content of various ethanol yeast products contained 10.3 mg/100 mg. The mannan content of DDGS was
1.0 mg/100 mg, and the mannan content of a new (50% crude protein), maize distillers dried yeast (AAFCO – 96.5) co-product was
3.0 mg/100 mg. Therefore, by calculating the percentage of mannan content of maize co-products relative to mannan content in pure
71
ethanol yeast products suggest that DDGS contains about 10% yeast and the high protein maize co-product contains about 29% yeast.
Lim et al. (2009) used an enzymatic method (AACC method 32–23.01) designed to measure β-glucan content in barley and oats,
and estimated that DDGS contained 0.57% β-glucans. However, it is not clear if this method is suitable for fermented grain co-
products. Kim et al. (2008) estimated that the total glucan concentration in DDGS is 21.2% (dry matter basis), of which 16% was
attributed to cellulose and 5.2% was from starch, using an average analysis of cellulosic biomass composition analysis from 3 research
laboratories. However, while glucans are found in bacteria, fungi, yeast, and cereal grains, they vary in molecular configuration,
solubility, and functionality. Cellulose is not considered a functional source of β-glucans because it is insoluble and does not provide
the same functionality of yeast β-glucans (Sikora et al., 2013). de Vries et al. (2016) used a method from Megazyme International Ltd.
(Ireland) to determine the β-glucan content and digestibility of an extract added to DDGS and rapeseed diets. Recent unpublished
data (T. Politano, United Wisconsin Grain Processors, LLC, Friesland, WI, personal communication) using the Megazyme Interna-
tional Yeast β-glucan assay showed that the β-glucan content in for a new 50% crude protein, maize distillers dried yeast (AAFCO –
96.5) co-product was 8.2–8.4% on an “as-fed” basis. Use of the Megazyme assay appears to provide good accuracy and minimal cross-
reactivity with starch. Until further studies are conducted to compare the accuracy of β-glucan assay methodologies for yeast con-
taining products, use of the Megazyme assay seems appropriate.
Alizadeh et al. (2016) used a gas-liquid chromatography procedure described by Englyst and Cummings (1988) with some
modifications (Slominski and Campbell, 1990) to determine the non-starch polysaccharide (including mannose) content of yeast-
derived products and a wheat-corn DDGS source. The concentration of mannose in wheat-corn DDGS was 1.6%. They also determined
the concentration of nucleotides to be 0.13% using an internal HPLC method developed by Canadian Bio-Systems Inc. (Calgary,
Alberta, Canada).
6. Conclusions
Numerous yeast supplements and yeast-containing feed ingredients are produced, marketed and used in animal feeds as nutrient
sources, probiotics, providers of nutraceutical compounds (i.e. β-glucans, mannanoligosaccharides, nucleotides), or serve unique
nutritional functions (i.e. Se yeast and Phaffia yeast). A large number of scientific studies have generally shown that yeast and its
derivatives, may provide beneficial effects on animal growth performance and health, especially when animals are reared in sub-
optimal hygienic conditions or are experiencing a disease challenge. It is essential that nutritionists understand the inherent dif-
ferences between these products, along with their potential role in providing desired potential effects when added to animal feeds.
While numerous analytical methods have been developed and compared for determining viable yeast content and yeast viability in
active dried yeast products, each method has limitations, and there is no standard procedure being used by the U.S. or global feed
industry. Even more concerning is the lack of suitable, commercially available analytical methods to determine dead yeast and yeast
cell components in yeast containing additives and ingredients. However, the use of mannan content of pure yeast products as a proxy
for estimating yeast content in maize co-products appears to be more accurate than using amino acid profiles, and the use of the
commercially available Megazyme International Yeast β-glucan assay to determine β-glucan content in maize co-products appears to
be an accurate and practical method. Appropriate analytical methodologies are necessary to assess functional nutritional components
in yeast-containing products to help nutritionists quantify these components, and use them strategically in precision animal nutrition
feeding programs.
Conflict of interest
I am the sole author of this manuscript and have no conflict of interest.
Acknowledgements
Special thanks to Klaus Englyst (Englyst Carbohydrates, Ltd., Southampton, UK) for providing analytical data for mannan content
of yeast containing products, Tim Politano (United Wisconsin Grain Processors, LLC, Friesland, WI) for providing analytical data for
β-glucan content in maize co-products, and Dr. Peter Williams, AG-BIO Ltd., Northampton, UK for helpful suggestions in writing this
review. Support for these contributions was provided by CHS, Inc. (Inver Grove Heights, MN, USA) and Fluid Quip Process
Technologies (Cedar Rapids, IA, USA).
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Animal Feed Science and Technology 235 (2018) 60–76

Contents lists available at ScienceDirect

Animal Feed Science and Technology

Yeast and yeast derivatives in feed additives and ingredients:

2. Yeast species of commercial importance

3. Yeast and yeast-containing feed ingredients

3.1. Viable yeast products

3.2. Yeast culture

3.3. Nutritional yeast products

Component, % Brewer’s yeast Dehulled soybean meal

Dry matter 93.30 ± 2.5 87.8 ± 0.6

3.4. Specialty yeast products

3.4.1. Irradiated yeast

3.4.2. Selenium yeast

3.4.3. Chromium yeast

3.4.4. Phaﬃa yeast

3.5. Fractionated yeast products

3.6. Yeast and yeast components in corn ethanol co-products

Animal species Yeast probiotic responses References

4.1. Probiotic eﬀects of live yeast

4.2. Health beneﬁts of yeast cell wall components

Body weight gain +1.75 (44) 35 6 3

Body weight gain +2.25 (27) 19 8 0

ADG2 +4.12 (54) 36 17 1

Parameter No. Studies Reporting an Improvement No. Studies Reporting No Change

Immune response Reference

No eﬀects Hiss and Sauerwein (2003)

No change Increase Decrease No change Increase Decrease No change Increase Decrease

4.2.5. Mycotoxin binding

4.2.6. Immune responses from feeding DDGS

5. Methods to measure yeast and yeast derivative concentrations

5.1. Direct measurement of yeast cells

5.2. Methods to determine yeast viability and vitality

5.3. Methods to determine yeast and yeast-derivatives in DDGS

I am the sole author of this manuscript and have no conﬂict of interest.

of astaxanthin-rich Phaffia rhodozyma in feedingstuffs for salmon and trout. . https://ec.europa.eu/food/sites/food/files/safety/docs/animal-feed_additives_rules_

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