Letters in Applied Microbiology ISSN 0266-8254

ORIGINAL ARTICLE

The health enhancer yeast Saccharomyces cerevisiae in two

types of commercial products for animal nutrition
J.F. Garcia-Mazcorro1,2 , M.V. Rodriguez-Herrera3, A.G. Marroquin-Cardona4 and J.R. Kawas5
1 Research and Development, MNA de M
exico, San Nicola
s de los Garza, M

exico
2
noma de Nuevo Leo
Faculty of Veterinary Medicine, Universidad Auto
n, General Escobedo, M
exico
3
n e Innovacio
RT-Biotech, Parque de Investigacio
n Tecnolo

gica, Apodaca, M
exico
4
noma de Nuevo Leo
Department of Physiology, Pharmacology and Toxicology, Faculty of Veterinary Medicine, Universidad Auto
n, General
Escobedo, M
exico
5 Faculty of Agronomy, Universidad Auto
noma de Nuevo Leo
n, General Escobedo, M
exico

Significance and Impact of the Study: Probiotics (or direct fed microbials) are increasingly popular in
Animal Nutrition. Different products containing live micro-organisms or microbial-derived products are
commercially available to enhance health and boost commercial traits. The characteristics of these prod-
ucts dictate their physiological effects and determine their potential to increase profitability from live-
stock. For the first time, this report presents data about the numbers and phenotype of the health
enhancer Saccharomyces cerevisiae in two widely available commercial products in Animal Nutrition.
These findings may be useful for scientists and producers around the globe and have the potential to
open up novel venues for research.

Keywords
antibiotics, fermentation biotechnology,
mechanism of action, probiotics, yeasts.
Correspondence
Jose F. Garcia-Mazcorro, Department of
Research and Development, MNA de Mexico,
Avenida Acapulco 770, Col. La Fe, San
Nicolas de los Garza, Nuevo Leon 66477,
Mexico.
E-mail: josegarcia_mex@hotmail.com
2019/0009: received 13 November 2018,
revised 12 February 2019 and accepted 20
February 2019
doi:10.1111/lam.13141
Abstract
The health enhancer yeast Saccharomyces cerevisiae (SC) is widely used in diets
for different animals. Two main types of SC-based products are commercially
available, one containing live yeasts and one containing SC fermentation by-
products, which are supposedly not dependent on live yeasts for their
physiological effects in vivo. Culture-based techniques were applied to study
yeasts in two types of commercial products: a product containing live SC
(LSC) and a SC fermentation product (SCFP). Three temperatures (25, 30 and
39°C) and two pH levels (4 and 7) were tested. The product with LSC
contained an average of 1
21 9 109 colony-forming units (CFUs) of yeasts per
g contents (min: 1 9 108, max: 3 9 109). In contrast, the SCFP contained an
average of 4
67 9 103 (min: 3 9 102, max: 1
9 9 104) CFUs per g contents (c.
1 million times less than the concentration of yeasts in the product with LSC).
Both temperature and pH level affected the number of CFUs but this effect
differed between the two products. Biochemical tests identified the two yeasts
as SC, which differed in their ability to ferment maltose (negative in the
SCFP). This report encourages more research on commercial microbial strains
for animal nutrition that can lead to a better understanding of their mode of
action in vivo.

animal feed and dozens of these products are commer-

Introduction
Probiotics are live micro-organisms that, when adminis-
tered in adequate amounts, confer a health benefit on the
host (FAO, WHO 2002). The Food and Drug Administra-
tion (FDA) of the USA uses the name direct-fed micro-
bials (DFM) for products containing probiotics used in
cially available for animal nutrition (FAO 2016). DFM
contain various numbers of one or more species of bacte-
ria or yeasts and growing evidence suggest they can be
used as an effective alternative to Antibiotic Growth Pro-
moters (FAO 2016). Despite the inherent wide variability
among the different studies, there is enough evidence to

472 Letters in Applied Microbiology 68, 472--478 © 2019 The Society for Applied Microbiology
J.F. Garcia-Mazcorro et al.
suggest that some DFM can improve health and produc-
tivity in animals under some circumstances (Chau-
cheyras-Durand et al. 2008; Buntyn et al. 2016).
Yeast products have long been used as DFM in diets
for different animal species (particularly livestock species)
to improve health and productivity (Osmakova et al.
1964; Chaucheyras-Durand et al. 2008; Uyeno et al.
2015). The most commonly used yeast for this purpose is
Saccharomyces cerevisiae (SC). Importantly, most papers
have focused on the effects of SC on animal health and
productivity in vivo and much fewer papers have focused
on the mode of action of the micro-organisms (Gray and
Ryan 1990; Newbold et al. 1996; Fonty and Chaucheyras-
Durand 2006; Marden et al. 2008), in part due to the
complexity associated with this phenomenon. For
instance and despite the recent massive efforts to study
the gut microbiota of ruminants and other animal spe-
cies, to our knowledge no one has shown direct proof
that SC (a ‘typical aerobe’, accordingly to the American
Type Culture Collection, ATCC) can reproduce inside the
gastrointestinal tract of livestock, although this seems to
be unlikely based on in vitro studies (Graff et al. 2008)
and studies in humans, mice and other animals (Klein
et al. 1993; Edwards-Ingram et al. 2007; Garcia-Mazcorro
et al. 2011; Samonis et al. 2011). However, SC is one of
the few yeasts that can grow rapidly under anaerobic con-
ditions (Ishtar Snoek and Yde Steensma 2007).
There are two main types of commercial SC-based
products used in livestock operations. First, there are
commercial products containing live yeasts, for example
Yea-Sacc (Alltech, Nicholasville, KY), a well-known and
well-studied product. The European Food Safety Author-
ity published a comprehensive review on the safety and
efficacy of Yea-Sacc as a feed additive for several livestock
species, with doses varying from 5 9 107 to 1 9 108 col-
ony-forming units (CFUs) per kg of complete feed
depending on the animal species and phase of production
(EFSA Feedap Panel 2014). Importantly, the EFSA report
suggested a minimum dose of 5 9 107 CFUs per kg com-
plete feed (5 9 104 CFUs per g complete feed) to reach a
significant effect on milk production in cows. As a com-
parison, commercial products containing another type of
Saccharomyces used mostly in humans (S. boulardii) con-
tain anywhere from 5
8 logs (c. 6
3 9 105) to 10
1 logs
(c. 1
1 9 1010) CFUs per g content of one capsule/sachet
(usual daily dose), each weighting on average 357 mg
(Vanhee et al. 2010). S. boulardii has been called the pro-
biotic strain of SC (Edwards-Ingram et al. 2007) and is
different from other Saccharomyces in several key charac-
teristics (e.g. cell wall thickness) but shares >99% geno-
mic relatedness with SC and the genes associated with
probiotic properties in S. boulardii are also conserved in
SC (Khatri et al. 2017). Please note that the effect of live
Letters in Applied Microbiology 68, 472--478 © 2019 The Society for Applied Microbiology
Commercial yeasts for animal nutrition
yeast is thought to be quite different compared to the
effect of yeast fermentation products, and mainly refers to
the respiratory activity of the organisms (Gray and Ryan
1990; Newbold et al. 1996).
The other type of commercial SC-based products sup-
posedly does not contain live organisms (thus by defini-
tion cannot be catalogued as probiotics) and are generally
known as yeast fermentation products in the literature
(Zhu et al. 2017). These yeast cultures produced through
yeast fermentation contain fermentation by-products that
are not dependent on live yeast for their physiological
effects. Fermentation products may affect several in vivo
parameters through an altered ruminal microbial fermen-
tation by increasing fibre digestion, stimulating the
growth and activity of fibre-digesting and lactate-utilizing
bacteria, increasing microbial protein synthesis, and
decreasing accumulation of lactate (Zhu et al. 2017).
Please be aware of the controversy in the literature and
also in the market regarding these products (e.g. some
commercial yeast culture products are mislabelled as
products containing live yeasts, see comment in Poppy
et al. 2012, referring to the meta-analysis performed by
Desnoyers et al. 2009). The purpose of this investigation
was to evaluate the numbers and phenotype of yeasts
contained in commercial products commonly used in ani-
mal nutrition, one containing live SC and one containing
SC fermentation products (SCFP). The results may shed
light into the potential and variation in efficacy of these
products to improve health and productivity in vivo.
Results and discussion
The morphology of the yeast cells obtained directly from
the raw product containing live SC (i.e. without growing
them in Petri dishes) and the cells growing in the Petri
dishes from both products was visualized using light
microscopy (Fig. 1). Several biochemical tests identified
the two yeasts as SC and identical to each other with the
exception of maltose fermentation (negative in the fer-
mentation product, Table S1), despite being both positive
for maltose uptake.
A total of 36 Petri dishes were analysed for each pro-
duct (two samples, three temperatures, two pH levels, by
triplicate). The presence of live yeasts in the product with
live SC was detected with an average of 1
21 9 109 (min:
1 9 108, max: 3 9 109) CFUs per g of content (Fig. 2b).
To our knowledge, this is the first time that independent
scientists confirm the minimum amount of 1 9 108 CFUs
mentioned in the label of Yea-Sacc. There was no signifi-
cant difference between the two samples from the package
(P = 0
7167). There was a significant effect of both tem-
perature and pH level as well as Temperature*pH level
interaction (P < 0
0001 for all, please note that a
473
Commercial yeasts for animal nutrition J.F. Garcia-Mazcorro et al.

(a) (b)

L = 50·00 um L = 50·00 um

(c) (d)

L = 50·00 um L = 50·00 um

Figure 1 Saccharomyces cerevisiae (SC) cells. (a) Cells in the raw product containing live SC; (b) cells from the colonies of the product with live
SC; (c) cells from the colonies of the SC fermentation product; (d) cells from the colonies of a beer strain of SC (included here only for compara-
tive purposes). Scale bar = 50 lm. [Colour figure can be viewed at wileyonlinelibrary.com]

significant interaction implies that both factors together
played a role in the response variable, in this case the
number of CFUs). Multiple comparisons revealed inter-
esting differences or lack thereof among the different con-
ditions, for instance there was no difference in the
number of CFUs between 25 and 30°C for both pH levels
(Table 1) and there was no difference in the number of
CFUs between the two pH levels for both of these tem-
peratures, only at 39°C (Table 2).
The presence of live yeasts in the SCFP was detected
with an average of 4
67 9 103 (min: 3 9 102, max:
1
9 9 104) CFUs of yeasts per g of content (Fig. 2a).
There was no significant difference between the two sam-
ples from the package (P = 0
9287) and there was no sig-
nificant effect of pH level (P = 0
1730). However,
Temperature and the Temperature*pH level interaction
were significant (P < 0
001). Multiple comparisons
revealed interesting differences among the different
conditions that did not necessarily resemble the results
obtained from the product containing live SC (Tables 1
and 2). For instance while the numbers of CFUs in the
product with live SC were consistently higher at pH 7,
the numbers of CFUs in the SCFP showed an opposite
trend of growth at different pH levels depending on the
temperature (Fig. 2a).
The minimum concentration of live micro-organisms
(either bacteria or yeasts) that is capable of inducing a
biologically significant effect in the live host is a matter of
debate. In human medicine, Hill et al. (2014) suggested a
minimum level of 1 9 109 CFU per serving to be consid-
ered as probiotics but one dose is unlikely to produce any
measurable beneficial effect; in other words, the consump-
tion of probiotics must be continuous for at least a few
days in order to have a beneficial effect. The numbers
suggested by Hill et al. are particularly interesting in the
case of Saccharomyces. As mentioned above, commercial
products containing S. boulardii contain anywhere from
5
8 logs (c. 6
3 9 105) to 10
1 logs (c. 1
1 9 1010) CFUs

474 Letters in Applied Microbiology 68, 472--478 © 2019 The Society for Applied Microbiology
J.F. Garcia-Mazcorro et al. Commercial yeasts for animal nutrition

(a) (b) 9·5

4·0 9·3

CFUs/g product
9·0
CFUs/g product

3·5
8·8

8·5
3·0
8·3

2·5 8·0
25 30 39 25 30 39
Temperature Temperature

Figure 2 Box plots showing colony forming units (CFUs, in logs) per gram of product. (a) Saccharomyces cerevisiae (SC) fermentation product;
(b) product with live SC. Lines represent medians, plus symbols represent means, circles represent outliers (i.e. points that are a distance of more
than 1
5 times the intra-quartile range from the box, accordingly to SAS). See Tables 1 and 2 for statistical results. pH.level ( ) 4, ( ) 7. [Colour
figure can be viewed at wileyonlinelibrary.com]

Table 1 Summary of statistical results from the comparison of logs of Table 2 Summary of statistical results from the comparison of logs
colony-forming units among different temperatures at two pH levels of colony-forming units between the two pH levels at different
temperatures
pH
level 25°C vs 30°C 25°C vs 39°C 30°C vs 39°C Product with live SC SC fermentation product

Product with live Saccharomyces cerevisiae (SC) pH 4 vs pH 7

pH 4 Difference* (CL†): Difference (CL): Difference (CL): 25°C Difference* (CL†): Difference (CL):
0
32 ( 0
19/0
84) 2
37 (1
85/2
88) 2
04 (1
53/2
55) 0
20 ( 0
71/0
31) 0
20 ( 2
32/ 0
08)
P = 0
4015 P < 0
0001 P < 0
0001 P = 0
8341 P = 0
0298
pH 7 Difference (CL): Difference (CL): Difference (CL): 30°C Difference (CL): Difference (CL):
0
41 ( 0
10/0
92) 1
36 (0
85/1
87) 0
95 (0
44/1
46) 0
12 ( 0
63/0
39) 1
43 (0
31/2
55)
P = 0
1796 P < 0
0001 P < 0
0001 P = 0
9800 P = 0
0067
SC fermentation product 39°C Difference (CL): Difference (CL):
pH 4 Difference (CL): Difference (CL): Difference (CL): 1
21 ( 1
72/ 0
69) 0
66 ( 0
46/1
78)
0
73 0
99 ( 0
12/2
12) 1
72 (0
61/2
85) P < 0
0001 P = 0
4750
( 1
84/0
39) P = 0
1009 P = 0
0008
P = 0
3751 P values were adjusted by the Tukey’s method in SAS. Significant
pH 7 Difference (CL): Difference (CL): Difference (CL): comparisons (P < 0
05) are highlighted for better visualization.
1
90 2
87 0
96 SC, Saccharomyces cerevisiae.
(0
78/3
02) (1
75/3
99) ( 0
15/2
08) *Difference implies differences in average values.
P = 0
0002 P < 0
0001 P = 0
0808 †CL implies 95% confidence limits (i.e. the difference between the
means is 95% likely to fall within this range).
P values were adjusted by the Tukey’s method in SAS. Significant
comparisons (P < 0
05) are highlighted for better visualization.
*Difference implies differences in average values.
†CL implies 95% confidence limits (i.e. the difference between the
suggested amount (5 9 104 CFUs per g complete feed) of
means is 95% likely to fall within this range). Yea-Sacc to increase milk production in dairy cattle (EFSA
Feedap Panel 2014). In other words, the numbers of live
yeasts in the SCFP are likely not to exert an effect by
per g content of one capsule/sachet (usual daily dose), themselves (aside from fermentation products). However,
each weighing on average 357 mg (Vanhee et al. 2010). please note that the recommended dose of the SCFP in the
Regardless of the original ingested number of micro-organ- product’s label is often several times higher compared to
isms, not all of the ingested probiotics survive the harsh the recommended dose of Yea-Sacc, depending on the ani-
conditions of the GI tract and they are rapidly eliminated mal species and the phase of production (Table 3), and
from the digestive tract. In this regard, our results raise the survival rate and reproductive capacity of the yeasts in
new intriguing questions. For instance the highest counts commercial products has not been investigated, something
of live yeasts in the SCFP (almost 2 9 104 CFUs per g of that may also have an impact on the ingested dose
product) are much lower compared to the minimum acquired by individual animals.

Letters in Applied Microbiology 68, 472--478 © 2019 The Society for Applied Microbiology 475
Commercial yeasts for animal nutrition J.F. Garcia-Mazcorro et al.

Table 3 Recommended doses in the products labels but it is possible that such small amounts of yeasts are
not relevant for the company. Importantly, SC can enter
Animal species/
phase of production Dose Yea-Sacc Dose Original XP stationary phase (Hartwell 1974), alter their morphology
to a filamentous form (Gimeno et al. 1992), or sporulate
Beef cattle (Freese et al. 1982; Knight and Goddard 2016) and here
All NA 20–45 g per head per
we did not investigate the specific stage of the live yeasts
day (direct addition)
3–6 kg per ton feed
in the products. It is also possible that the SCFP studied
(concentrate feed) here (and perhaps other similar commercial products)
Initiation 2 kg per ton feed NA may contain environmental SC strains (B€ uchl et al. 2010),
Growth 1 kg per ton feed NA which, if true, would be interesting for biomedical scien-
Ending 0
5 kg per ton feed NA tists because of the potential mixture of various pheno-
Dairy cattle types (including survival and reproduction capabilities).
All NA 30–60 g per head per
Please note that our results from the SCFP (Fig. 2a) show
day (direct addition)
3–6 kg per ton feed
a wide variation in CFUs counts and a differential beha-
(concentrate feed) viour at different pH levels and temperatures, thus
Initiation 2 kg per ton feed NA strongly suggesting that the product contains more than
Growth 1 kg per ton feed NA one strain of SC. More research is needed (and ongoing
Lactation 1 kg per ton feed NA in our research facilities) in this regard.
Pigs Here we performed a few biochemical tests and noticed
Pre-initiation/ 2 kg per ton feed 8 kg per ton feed
that the two yeasts differed in maltose fermentation, and
initiation
Initiation/growth 0
5 kg per ton feed 8 kg per ton feed
reasoned that it was important to comment about this.
Ending 0
25 kg per ton feed NA Maltose fermentation in SC requires a set of MAL genes
Gestation 1 kg per ton feed 8 kg per ton feed (i.e. genes encoding a maltose transporter (MALT), mal-
Pre-partum/ 2 kg per ton feed 8 kg per ton feed tase (MALS), and zinc finger-type transcription factors
lactation (MALR). These genes are vital for fermentation of mal-
Chickens tose as demonstrated by the observation that yeasts lack-
Broiler chickens 1 kg per ton feed 3–5 kg per ton feed
ing any of these three genes fail to grow in medium with
Laying hens 1 kg per ton feed 3–5 kg per ton feed
Turkeys 1 kg per ton feed NA
maltose as the sole carbon source (Goldenthal et al.
Horses 1 kg per ton feed 30–60 g per head per 1987). In a medium containing high concentrations of
(adults/foals) day (direct addition) glucose, maltose uptake is transcriptionally repressed, a
4–6 kg per ton feed phenomenon that is known as glucose repression. Duan
(concentrate feed) et al. (2018) recently showed that wild and domesticated
Rabbits 2 kg per ton feed NA SC strains differ in their ability to utilize maltose. Inter-
Dogs and cats 2–3 kg per ton feed NA
estingly, the ‘domesticated’ milk SC isolates from tradi-
Please note that these suggested doses are interesting since there are tionally fermented dairy products from remote pastoral
no studies for some of these species (e.g. Yea-Sacc in horses, dogs or areas from western and northern China and Mongolia
cats). Also, this information was obtained from products sold in also showed high maltose utilization ability, even though
Northern Mexico and we do not know whether these indications are maltose is absent in milk. Clearly more research is needed
the same in other regions of the world.
into the ability of commercial SC strains to utilize mal-
NA, not specified in the product label.
tose and other substrates and the consequences of this
phenomenon for in vivo performance of livestock.
This paper shows that a commercial SCFP contains Our results have important implications for nutrition
viable yeasts and that these yeasts are SC. Interestingly, companies aiming to develop new Saccharomyces-based
Original XP and the other two SCFPs from Diamond V products. For instance the cost of production of products
(Original XPC and Original XPC ULTRA, see http:// containing live yeasts is substantially higher compared to
www.diamondv.com/products/) do not specify that they the cost of production of fermentation products, a differ-
contain active, live or viable organisms. Other products ence that is also reflected in the customer’s prices. Inter-
from the company such as the XP DFM do specify that estingly, the two products studied here are sold in the
they contain live micro-organisms (eight species of bacte- same presentation (i.e. stitched 25 kg sacks) and have sim-
ria plus the fungus Aspergillus oryzae) but does not specify ilar expiration dates, despite the fact that the contents are
that they contain live SC (http://www.diamondv.com). quite different, both in nutrient composition and (based
Overall we feel sceptical about the possibility that Dia- on this study) in the concentration of micro-organisms.
mond V is unaware that their product contain live yeasts Despite all limitations of this short communication, we

476 Letters in Applied Microbiology 68, 472--478 © 2019 The Society for Applied Microbiology
J.F. Garcia-Mazcorro et al.
believe that our findings can be useful for a wide variety
of people around the globe and have the potential to open
up new venues for novel research.
Materials and methods
Before designing a large study assessing lot and package
variation across different geographical regions and times
of year, here we decided to first focus on the numbers of
yeasts and their phenotype in commercial packages, simi-
larly to other studies investigating commercial formula-
tions containing S. boulardii for humans (Vanhee et al.
2010). One new 25 kg package of Yea-Sacc (Alltech, lot #
006983-2, contains a minimum of 1 9 108 CFUs per g of
live SC accordingly to the product label, ingredients based
on product’s label: SC grown on ground yellow corn, dia-
static malt and cane molasses to preserve the fermentative
activity) was opened up and two samples (200 g each,
from the top and one from the bottom section of the
package) were obtained to account for intra-package vari-
ability using (whirl-pak sampling bags, NY) and trans-
ported to the laboratory. Each sample was thoroughly
mixed inside the bag, 1 g was obtained, mixed with 9 ml
of auto-claved physiological saline (0
9% NaCl) and vari-
ous tenfold dilution series (100 ll in 900 ll of saline) were
made (Vanhee et al. 2010). Ultimately, the objective of the
dilutions was to have approx. 10–50 CFUs per Petri dish
for ease and accuracy of count. The liquid containing the
final dilution was then placed in petri dishes containing
autoclaved PDA agar and grown at 25 and 30°C (ATCC
guidelines for most strains) and 39°C (rumen tempera-
ture) for 72 h at a pH of 4 (best pH to grow SC accord-
ingly to Salari and Salari 2017) and 7 (upper limit of
rumen pH according to Van Kessel and Russell 1996). The
pH of PDA media was adjusted with 1 mol l 1 HCl and
2 N NaOH before sterilization (15 min at 121°C and 15
psi) and measurements of pH in agar plates stayed similar
at the beginning and the end of growth. Additionally, we
performed the exact same procedure with a 25 kg package
of the SCFP Original XP (Diamond V, Cedar Rapids, IA,
lot # A010418, ingredients based on product’s label: SC
grown on corn flour, corn gluten, wheat bran, rye bran,
and cane molasses). Therefore, a total of 36 samples (two
samples, three temperatures at two pH levels, three repli-
cates each) from each product (SCFP and product with
live SC) were used in this study. All samples were pro-
cessed within a week, and each set of conditions (e.g. 25°C
at pH 4) included samples from both products. Negative
controls (saline) were used in each batch of samples. Bio-
chemical testing was performed by the Laboratory of Clini-
cal Chemical Analysis at the Faculty of Chemical Sciences
of the Universidad Aut
onoma de Nuevo Le
on (UANL,
Monterrey, M
exico). The GLIMMIX procedure in SAS
Letters in Applied Microbiology 68, 472--478 © 2019 The Society for Applied Microbiology
Commercial yeasts for animal nutrition
University Edition was used to compare the log numbers
of CFUs among the different conditions. The residuals
from this analysis did not significantly deviate from a nor-
mal distribution (Fig. S1), thus suggesting that the model
was appropriate for our data (see more about this in
Supporting Information).
Acknowledgements
This study was partly supported by MNA de M
exico, a
company of Animal Nutrition.
Conflict of Interest
J.F.G.M. is an employee of MNA de M
exico, a company
of Animal Nutrition. M.V.R.H. is an employee of RT-Bio-
tech, a company that commercializes live yeast mainly for
beer production.
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Supporting Information
Additional Supporting Information may be found in the
online version of this article:
Figure S1 Residuals plots for the described models
(CFUs = Temperature pH.level Temperature*pH.level) in
GLIMMIX for Original XPTM (a) and Yea-Sacc (b).
Table S1 Results of biochemical reactions for the two
products.