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Microbial community structure and diversity in different
types of non-bovine milk
Wei Wei1, Xinyu Hu1, Zhaozhi Hou2, Yuan Wang2 and Lin Zhu2
Non-bovine milk is an underappreciated type of milk with
geographical characteristics of production, consumption and
nutrition. Although the structure similarity of microbial
community from the different types of non-bovine milk are very
different, the composition of microbial community, especially
those belonging to pathogenic bacteria or lactic acid bacteria in
non-bovine milk, may be more diverse than that of cow milk.
Microbial community analysis based on high-throughput
sequencing coupled with microbial annotation on species level,
environmental factors of milk producing areas, and milk
nutritional composition is considered to be very necessary
strategy in future research of non-bovine milk. This review
provides a reference for evaluating food safety and nutrition of
non-bovine milk and dairy products.
Addresses
1
School of Agricultural Engineering, Jiangsu University, Zhenjiang,
China
2 munity diversity, remains extremely limited [11,12].
School of Food and Biological Engineering, Jiangsu University,
Zhenjiang, China
Corresponding author: Wei, Wei (weiwei7096@ujs.edu.cn)
Current Opinion in Food Science 2021, 38:51–57
This review comes from a themed issue on Food microbiology
Edited by Anderson de Souza Sant’Ana
https://doi.org/10.1016/j.cofs.2021.01.008
2214-7993/ã 2021 Elsevier Ltd. All rights reserved.
Introduction
Milk is a highly nutritious food for human consumption
and can be obtained from a variety of animals, including
human [1,2]. Dairy cows and water buffalos account for
more than 90% of the global milk production (Figure 1a)
[3,4]. Therefore, people have devoted constant and close
attention to issues such as food safety and nutritional
components in milk [5,6]. In addition, non-bovine milk,
such as goat, donkey, camel, horse, and breast milk,
accounts for most of the remaining milk production.
Although the absolute yields of these non-bovine types
of milk are low, they have certain differences in
the geographical distribution, nutritional components,
and therapeutic uses. For example, because of its low
fat and high iron contents, goat milk has become a
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designated raw material for certain soft cheeses in the
Mediterranean region [7], where is the main producing
area of goat milk in the world (Figure 1a). Donkey milk is
used as a medicinal food in Italy, providing an important
solution for the treatment of young children affected by
milk allergies [4]. Camel milk is an almost exclusive
source of animal milk in Africa and the Arab region
(Figure 1a) [7,8]; horse milk is a special animal milk
for nomadic ethnic groups in China’s Inner Mongolia and
Xinjiang Uygur regions (Figure 1b) [9]; and breast milk
provides essential nutrients, immune cells, and bioactive
components for infants [7,10]. Thus, these non-bovine
types of milk play an important role in the field of human
nutrition, health and consumption in some geographical
region. However, research on these non-bovine types
of milk, which contribute to food safety and human
nutritional health, especially research on microbial com-
The rich variety of nutrients in the milk and its near-
neutral pH condition, which are similar to those of natural
basic media, provide favourable conditions for microbial
reproduction [1]. In previous studies of milk microbial
communities, researchers have used culture-free detec-
tion methods to obtain information on fungal internal
transcribed spacer fragments of Geotrichum, Malassezia,
Torrubiella, Trichosporon and Kluyveromyces; however,
these fungi mostly occur when animals are in poor health
or are maintained under poor feeding conditions [13,14].
Bacteria still dominate microbial communities in
the milk of normal and healthy animals. Therefore,
current microbial studies of non-bovine milk, irrespective
of whether they are based on culture-dependent or
culture-independent techniques, are almost exclusively
focused on bacterial communities [11].
Similarity in the bacterial community structure
in different types of non-bovine milk
Taking goat, donkey, camel, horse, and breast milk as the
target samples, we screened 126 reports about microbial
communities in non-bovine milk, involving 154 genera
belonging to 7 phyla of bacteria (Table S1). Among these
genera, 51 genera were obtained by culture-dependent
techniques and 152 genera were determined by culture-
independent techniques (Table S1 in Supplementary mate-
rial). Heatmap analysis of the occurrence of these bacterial
genera in all tested samples of non-bovine milk was
performed (Figure 2a). We found that more bacterial genera
were detected in cow, breast and goat milk samples
than those of other non-bovine milk samples under
Current Opinion in Food Science 2021, 40:51–57
52 Food microbiology

Figure 1

(a)

Cow milk Buffalo milk

Goat milk Camel milk

(b)

Horse milk Donkey milk Horse and donkey milk

Current Opinion in Food Science

Heatmap of production or consumption of different types of milk in the world (except Antarctica). (a) Schematic profiles of the production of cow,
buffalo, goat and camel milk. Data comes from Food and Agriculture Organization of United Nations of 2018 (http://www.fao.org/faostat/en/). The
heatmap value is calculated by the formula of log10T, in which T indicates the production of raw milk (Tonnes). (b) Schematic profiles of the
consumption of horse and donkey milk. Consumption reports are used instead of production data that cannot be obtained. If horse milk or donkey
milk is recorded to be consumed in this area, which is indicated by purple or orange, both are consumed in an area indicated by brown.

culture-dependent and culture-independent techniques,
especially culture-independent techniques (Figure 2a).
These results indicate that breast and goat milk may be
able to inhabit more diverse bacterial populations as well as
cow milk, and culture-independent technique is very neces-
sary for the deep exploration of such bacterial populations in
non-bovine milk. Therefore, to analyse the bacterial com-
munity structure and diversity in different types of non-
bovine milk, high-throughput sequencing results, targeting
bacterial 16S rRNA gene fragments, were screened.
High-throughput sequencing studies of bacterial commu-
nities in non-bovine milk, which were published after
2013, were collected, and the corresponding data for
47 non-bovine milk samples of five different types,
including goat, donkey, camel, horse, and breast milk,
were obtained (Table S2 in Supplementary material). In
addition, data on 10 cow milk samples from different
geographical regions were collected as controls (Table S2
in Supplementary material). Considering differences in
the used primers and sequencing platforms, the annota-
tion information for bacterial genera with relative abun-
dances of greater than 0.1% was directly extracted for all
samples. Based on the community composition and rela-
tive abundances of these dominant bacteria, principal
component analysis (PCA) of the structural similarity of

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 Microbial community in non-bovine milk Wei et al. 53

Figure 2

Culture-dependent Culture-independent
other phyla Horse milk, China, Xilin Gol Aera
(a) (b) Goat milk, China, Guanzhong Aera
Bacteroidetes
Goat milk, China, Jilin Province

Actinobacteria 5 Donkey milk, Italy, Northwest

Donkey milk, Italy, Martina Franca

Cow milk, Ireland
Cow milk, New Zealand, Northland and Southland

PC2 (8.9%)
Cow milk, China, Shanghai

Cow milk, Norway, southern part

Proteobacteria Cow milk, Italy, Trento
0
Camel milk, Algeria, Ghardaia
Camel milk, China, Xinjiang Proxince
Camel milk, China, Inner Mongolia
Breast milk, Netherlands, Nijmegen
Breast milk, SouthAfrica, CapeTown
Breast milk, Ireland, Cork

Firmicutes -5 Breast milk, Spain, Valencia

Breast milk, America, Washington State
Breast milk, Canada
Breast milk, China, Beijing
Breast milk, Finland, southwestern area

-5 0 5
H w

H w
C t

C t
D el

D el
an

an

se
y

y
oa

oa
ke

ke
o

o
am

am

or
um

um
C

C
G

G
on

on

PC1 (9.5%)
Current Opinion in Food Science

Diversity and similarity of bacterial communities in different types of non-bovine milk. (a) Heatmap of diverse bacterial population isolated and
detected from different types of non-bovine milk based on culture-dependent and culture-independent techniques, respectively. Green colour
indicates that bacterial genera has been reported in a type of cow and non-bovine milk, otherwise in light grey colour. Bacterial genera belonging
to the same phylum were clustered and represented by the same colour. Other phyla includes Deinococcus-Thermus, Fusobacteria and
Verrucomicrobia. (b) Principal component analysis (PCA) plot shows structure similarity of bacterial community in different non-bovine milk
samples based on the operational taxonomy units (OTUs) of bacterial 16S rRNA gene from the high-throughput sequencing data. Different symbol
shapes indicate different milk samples, and different colours of the same shape indicate milk samples in different sampling areas.

bacterial communities in different types of milk was
performed (Figure 2b).
The PCA results showed that the PCA scores of the
microbial community of breast milk samples from differ-
ent regions were uniquely distributed in the second
quadrant (Figure 2b), which indicated that the structure
similarity between the microbial community of breast
milk and that of other animal milk was low. Among all
the tested animal milk samples, goat, camel and cow milk
had more similar bacterial community structure, and
were different from those of horse and donkey milk
(Figure 2b), which indicated that there might be different
bacterial community structures in the milk of ruminants
and non-ruminants. These results suggested that the
similarity of microbial community structures from the
different types of non-bovine milk is very different.
It has been reported that the geographical distribution is
an important factor in the formation of the bacterial
community structure in cow milk [15]. However,
geographical distribution as an influence factor of milk
microbial community is too vague, and we believe that it
is more important and academic to substantialize the
influence factors caused by geographical distribution.
Studies have shown that microbes in animal milk mainly
originate from the producing environment of animal milk
[16]. Microbial sources of milk producing areas present
in different geographical environments may really
determine the diversity and functionality of microbial
communities in local non-bovine milk [17,18]. Therefore,
the environmental factors (such as temperature, precipi-
tation, light, humidity, etc.) affecting microbial commu-
nity in milk producing areas may be the substantial factors
determining the microbial community in non-bovine
milk [19].
Simultaneously, the milk is similar to a microbial selective
medium and can selectively enrich its surrounding envi-
ronmental microbes to form a specific milk bacterial
community [20]. Thus, the nutritional composition of milk
also plays a role in determining the microbial community
in non-bovine milk [21]. According to the previous
description, breast milk samples showed very different
bacterial community structures with those of animal milk,
regardless of whether they were collected in the same
region or in different regions (Figure 2b). These differ-
ences may be due to more diverse nutritional components
in breast milk than that in animal milk [18,22], which is
caused by the complex dietary composition and
dietary habits of humans [23]. These diverse nutritional
components are similar to different selective medium
formulations and can specifically select microbes and form
different microbial communities in breast milk.
In conclusion, the environmental factors of milk produc-
ing areas and milk nutritional composition may play
crucial role in the formation of microbial community in

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54 Food microbiology
non-bovine milk, which two aspects is just the lack of
research on microbial community in non-bovine milk.
Few report that combines the results of milk microbial
community, milk origin environmental factors and milk
nutritional composition. Therefore, we strongly suggest
that more attention should be paid to the effects of the
environmental factors of milk producing areas and milk
nutritional composition on milk microbial community in
the future research of non-bovine milk.
Species annotation of bacterial community
compositions in different types of non-bovine
milk
In milk bacterial communities, pathogenic bacterial spe-
cies are the primary concern because these bacteria cause
deterioration of milk and dairy products and seriously
endanger the human health [24,25]. With the increasing
demand for nutritional components and flavours of milk,
people begin to pay attention to the diversity and func-
tion of lactic acid bacteria (LAB) in milk [26,27]. In recent
years, using the high-throughput sequencing technology
based on nucleic acid sequences, animal milk has been
found to be the home for non-pathogenic and non-lactic
acid bacterial populations with a surprisingly high diver-
sity and relative abundance [28,29]. These microbial
communities non pathogenic and lactic acid bacteria also
play an important role in food safety, nutrition, and the
flavour of animal raw milk and dairy products. Therefore,
to further investigate the diverse bacterial community
composition in non-bovine milk, raw sequencing data of
the corresponding fragments of the bacterial 16S rRNA
gene sequence between primers 515F and 806R were
selected in the samples used by previous PCA for stan-
dardized analysis. After annotation of species and their
relative abundances in 46 different milk samples, 55 bac-
terial genera belonging to Proteobacteria, Firmicutes, Acti-
nobacteria, and Bacteroidetes, with relative abundances of
greater than 0.1%, were obtained, including 15 potentially
pathogenic bacterial genera, seven lactic acid bacterial
genera, and 33 non-pathogenic and non-lactic acid bacte-
rial genera (Figure 3).
In cow milk samples, two bacterial genera belonging to
Proteobacteria, namely, Pseudomonas and Acinetobacter, were
present, which are commonly present in almost all milk
samples. In non-bovine milk samples, bacterial genera
belonging to Proteobacteria had high relative abundances,
including 12 non-pathogenic genera and 11 potentially
pathogenic genera (Figure 3). At the same time, a notewor-
thy phenomenon was that within Proteobacteria, non-
pathogenic Betaproteobacteria had high relative abundances
in breast milk samples, while pathogenic Gammaproteobac-
teria had high relative abundances in other non-bovine milk
samples. Actinobacteria and Bacteroidetes exhibited low
relative abundances in all milk samples. Among them,
Actinobacteria only exhibited high diversity in breast milk
samples, whereas in most other non-bovine milk samples,
Current Opinion in Food Science 2021, 40:51–57
there was only one dominant and potential pathogenic
genus, Corynebacterium [30]. Bacteroidetes only showed
diversity in cow and donkey milk, with potentially patho-
genic Chryseobacterium and Flavobacterium as the dominant
bacterial genera. The results demonstrated that bacterial
communities in non-bovine milk samples had more diverse
populations, especially that in Proteobacteria, than those
of cow milk samples. Among the non-bovine milk,
breast milk was rich in diverse non-pathogenic bacterial
populations, while other types of non-bovine milk had
more potentially pathogenic bacterial populations, indicat-
ing that non-bovine milk of animal origin may have a
higher risk for food safety issues than breast milk and
cow milk.
All milk samples were rich in Firmicutes. Most of the other
genera aside from Staphylococcus, which may contain
pathogenic bacteria [31], were LABs (Figure 3). Among
these LABs, Lactococcus, Streptococcus, and Enterococcus
were the dominant genera in cow milk, while Streptococ-
cus, Lactococcus, and Lactobacillus were the dominant gen-
era in goat milk. Streptococcus was the dominant genus in
breast and camel milk, but its relative abundance in camel
milk was very low. Lactococcus, Leuconostoc, and Lactoba-
cillus were the dominant genera in donkey milk, and
Lactococcus was the absolute dominant genus in horse
milk. These results indicate that the LAB group in
non-bovine milk has a variety of genus composition. After
pasteurization, with significant decrease of thermolabile
LABs (e.g. Lactococcus) [32], donkey and horse milk may
lack LAB populations, while cow, goat, and camel milk
retain bacterial populations similar to that of breast milk,
namely, with the predominance of heat-resistant
Streptococcus thermophilus [33]. These results indicate that
drinkable goat and camel milk may still have a dominant
lactic acid bacterial species, S. thermophilus, after pasteur-
ization treatment, which is similar to breast milk. When
using pasteurized donkey and horse milk to prepare
corresponding dairy products, large amounts of LABs
are needed to compensate for the loss of characteristics
and the flavour of fermented dairy products.
Recommendations for microbial community
research in non-bovine milk
The above analysis shows large structural differences and
composition diversity of bacterial communities in non-
bovine and cow milk samples, among which non-bovine
milk samples have a high relative abundance and diver-
sity of potentially pathogenic bacterial populations. We
advocate that research strategies for microbial communi-
ties in non-bovine milk should be improved by introduc-
ing not only geographical and climatic conditions and
other environmental factors but also the basic physical
and chemical properties of milk in high-throughput
sequencing studies [19,34]. Furthermore, the structural
diversity of microbial communities in non-bovine milk
and influencing factors should be analysed using a variety
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 Microbial community in non-bovine milk Wei et al. 55

Figure 3

Current Opinion in Food Science

Species annotation, microbial type, relative abundance and phylogenetic analysis of dominant bacterial populations in 57 different cow and non-
bovine milk samples based on the high-throughput sequencing data of bacterial 16S rRNA gene. Symbols on tree tips specify major taxonomic
groups, including blue squares indicate Bacteroidetes, pink squares indicate Actinobacteria, green squares indicate Firmicutes, and purple
squares indicate Proteobacteria. The colour of the genus name is green to indicate the genus of lactic acid bacteria, and the genus name is red to
indicate that the genus may include pathogenic bacteria or opportunistic pathogens. Among the symbols representing different samples, different
shapes and colours respectively indicate different milk types and sampling areas. The relative abundance of bacterial genus is represented by a
circular symbol, and different colours and area of circles indicate different types of milk samples and relative abundance of bacterial genus,
respectively. The circle area is calculated by the formula of S = l g(x  10), in which x indicates the percentage value of relative abundance of
bacterial genus.

of bioinformatics methods to provide a powerful
reference for maintaining the quality and flavour of
non-bovine milk by regulating microbial communities.
In addition, the most recent sequencing platforms with
long-read sequencers, such as PacBio single-molecule
real-time (SMRT) sequencing, should be considered.
Compared with the currently widely used polymerase
chain reaction amplification and high-throughput
sequencing platforms based on 500-bp fragments (the
longest read), these new sequencing platforms can
complete full-length high-throughput sequencing of
the bacterial 16S rRNA gene, which can improve the
annotation of a bacterial community from the genus to the
species level [35,36,37]. This approach is very helpful
for more accurate identification of pathogenic bacterial
species and avoiding excessive or insufficient analysis of
bacterial species and their relative abundances in current
genus-level studies. We believe that the combination of
the above two strategies could allow more accurate anal-
ysis of microbial communities in non-bovine milk, con-
tribute to food safety and human nutritional health, and
provide an important reference for ensuring food safety

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56 Food microbiology
and the nutritional flavour of non-bovine milk and dairy
products.
Funding
This work was financially supported by the National Natural
Science Foundation of China (31770543, 31600069, 41503
068), Priority Academic Program Development of Jiangsu
Higher Education InstitutionsPAPD-2018-87), and Initial
Scientific Research Fund of Senior Talents in Jiangsu
University (15JDG016, 15JDG018).
Conflict of interest statement
Nothing declared.
Appendix A. Supplementary data
Supplementary material related to this article can be
found, in the online version, at doi:https://doi.org/10.
1016/j.cofs.2021.01.008.
References and recommended reading
Papers of particular interest, published within the period of review,
have been highlighted as:
 of special interest
 of outstanding interest
1. Quigley L, O’Sullivan O, Stanton C, Beresford TP, Ross RP,
Fitzgerald GF, Cotter PD: The complex microbiota of raw milk.
FEMS Microbiol Rev 2013, 37:664-698.
2. Astolfi ML, Marconi E, Protano C, Canepari S: Comparative
elemental analysis of dairy milk and plant-based milk
alternatives. Food Control 2020, 116:107327.
3. Balthazar CF, Pimentel TC, Ferrão LL, Almada CN, Santillo A,
Albenzio M, Mollakhalili N, Mortazavian AM, Nascimento JS,
Silva MC: Sheep milk: physicochemical characteristics and
relevance for functional food development. Compr Rev Food
Sci Food Saf 2017, 16:247-262.
4. Conte F, Panebianco A: Potential hazards associated with raw
 donkey milk consumption: a review. Int J Food Sci 2019,
2019:5782974
This review article summarizes the main microbial hazards possibly
present in raw donkey milk, and suggests that the microbial risk assess-
ment should performed along the whole milk chain. This review describes
diverse pathogenic microorganisms in different types of milk, which is a
helpful reference for the study of pathogenic microorganisms in non-
bovine milk.
5. Montgomery H, Haughey SA, Elliott CT: Recent food safety and
fraud issues within the dairy supply chain (2015–2019). Glob
Food Secur 2020, 26:100447.
6. Zwierzchowski G, Ametaj BN: Mineral elements in the raw milk
of several dairy farms in the province of Alberta. Foods 2019, 8.
7. Issa AT, Tahergorabi R: Chapter 22 - milk bacteria and
 gastrointestinal tract: microbial composition of milk. In
Dietary Interventions in Gastrointestinal Diseases. Edited by
Watson RR, Preedy VRBT-DI in GD. Academic Press; 2019:265-
275
This chapter reviews the sources of microbial contamination in different
types of milk and storage conditions for such microbial contamination.
Additionally, the diversity and function of lactic acid bacteria, pathogenic
bacteria and fungi in different types of milk were summarized. This is a
valuable reference for understanding the microbial diversity of milk.
8. Kashaninejad M, Razavi SMA: Influence of thermosonication
treatment on the average size of fat globules, emulsion
stability, rheological properties and color of camel milk cream.
LWT 2020, 132:109852.
Current Opinion in Food Science 2021, 40:51–57
9. Zhao F-Y, Liu Y-H, Zhi L, Li W-C, Xu H-Y, Sun Z-H, Bilige M,
Zhang H-P: Bacterial diversity of fresh mare’s milk in Xilingol.
Microbiol China 2019, 46:3295-3304.
10. Enjapoori AK, Kukuljan S, Dwyer KM, Sharp JA: In vivo
endogenous proteolysis yielding beta-casein derived
bioactive beta-casomorphin peptides in human breast milk for
infant nutrition. Nutrition 2019, 57:259-267.
11. McInnis EA, Kalanetra KM, Mills DA, Maga EA: Analysis of raw
goat milk microbiota: impact of stage of lactation and
lysozyme on microbial diversity. Food Microbiol 2015,
46:121-131.
12. Togo A, Dufour J-C, Lagier J-C, Dubourg G, Raoult D, Million M:
Repertoire of human breast and milk microbiota: a systematic
review. Future Microbiol 2019, 14:623-641.
13. Delavenne E, Mounier J, Asmani K, Jany J-L, Barbier G, Le Blay G:
Fungal diversity in cow, goat and ewe milk. Int J Food Microbiol
2011, 151:247-251.
14. Amrouche T, Mounier J, Pawtowski A, Thomas F, Picot A:
Microbiota associated with dromedary camel milk from
Algerian Sahara. Curr Microbiol 2020, 77:24-31.
15. Garroni E, Doulgeraki AI, Pavli F, Spiteri D, Valdramidis VP:
Characterization of indigenous lactic acid bacteria in cow milk
of the Maltese islands: a geographical and seasonal
assessment. Microorganisms 2020, 8.
16. Vacheyrou M, Normand A-C, Guyot P, Cassagne C, Piarroux R,
Bouton Y: Cultivable microbial communities in raw cow milk
and potential transfers from stables of sixteen French farms.
Int J Food Microbiol 2011, 146:253-262.
17. Tormo H, Ali Haimoud Lekhal D, Roques C: Phenotypic and
genotypic characterization of lactic acid bacteria isolated
from raw goat milk and effect of farming practices on the
dominant species of lactic acid bacteria. Int J Food Microbiol
2015, 210:9-15.
18. Kumar H, du Toit E, Kulkarni A, Aakko J, Linderborg KM, Zhang Y,
Nicol MP, Isolauri E, Yang B, Collado MC et al.: Distinct patterns
in human milk microbiota and fatty acid profiles across
specific geographic locations. Front Microbiol 2016, 7:1619.
19. Moossavi S, Sepehri S, Robertson B, Bode L, Goruk S, Field CJ,
 Lix LM, de Souza RJ, Becker AB, Mandhane PJ et al.:
Composition and variation of the human milk microbiota are
influenced by maternal and early-life factors. Cell Host Microbe
2019, 25:324-335.e4
This article illustrates that milk microbiota composition and diversity were
associated with maternal factors, breastfeeding practices, and other milk
components, showing an example of association analysis between milk
microbiota and the environmental factors.
20. Aakko J, Kumar H, Rautava S, Wise A, Autran C, Bode L, Isolauri E,
Salminen S: Human milk oligosaccharide categories define the
microbiota composition in human colostrum. Benef Microbes
2017, 8:563-567.
21. Li Z, Wright A-DG, Yang Y, Si H, Li G: Unique bacteria community
composition and co-occurrence in the milk of different
ruminants. Sci Rep 2017, 7:40950.
22. Padilha M, Danneskiold-Samsøe NB, Brejnrod A, Hoffmann C,
Cabral VP, Iaucci JD, Sales CH, Fisberg RM, Cortez RV, Brix S
et al.: The human milk microbiota is modulated by maternal
diet. Microorganisms 2019, 7:502.
23. Drago L, Toscano M, De Grandi R, Grossi E, Padovani EM,
Peroni DG: Microbiota network and mathematic microbe
mutualism in colostrum and mature milk collected in two
different geographic areas: Italy versus Burundi. ISME J 2017,
11:875-884.
24. Costanzo N, Ceniti C, Santoro A, Clausi MT, Casalinuovo F:
Foodborne pathogen assessment in raw milk cheeses. Int J
Food Sci 2020, 2020:3616713.
25. Cruzado-Bravo MLM, Barancelli GV, Dini Andreote AP, Saldaña E,
Vidal-Veuthey B, Collado L, Contreras-Castillo CJ: Occurrence of
Arcobacter spp. in Brazilian Minas frescal cheese and raw cow
milk and its association with microbiological and
physicochemical parameters. Food Control 2020, 109:106904.
www.sciencedirect.com

26. Reuben RC, Roy PC, Sarkar SL, Rubayet Ul Alam ASM, Jahid IK:
Characterization and evaluation of lactic acid bacteria from
indigenous raw milk for potential probiotic properties. J Dairy
Sci 2020, 103:1223-1237.
27. Valenzuela JA, Flórez AB, Vázquez L, Vasek OM, Mayo B:
Production of -aminobutyric acid (GABA) by lactic acid
bacteria strains isolated from traditional, starter-free dairy
products made of raw milk. Benef Microbes 2019, 10.
28. Li N, Wang Y, You C, Ren J, Chen W, Zheng H, Liu Z: Variation in
raw milk microbiota throughout 12 months and the impact of
weather conditions. Sci Rep 2018, 8:2371.
29. Zhao J, Fan H, Kwok L-Y, Guo F, Ji R, Ya M, Chen Y: Analyses of
physicochemical properties, bacterial microbiota, and lactic
acid bacteria of fresh camel milk collected in Inner Mongolia. J
Dairy Sci 2020, 103:106-116.
30. Oliveira A, Oliveira LC, Aburjaile F, Benevides L, Tiwari S,
Jamal SB, Silva A, Figueiredo HCP, Ghosh P, Portela RW et al.:
Insight of genus Corynebacterium: ascertaining the role of
pathogenic and non-pathogenic species. Front Microbiol 2017,
8:1937.
31. Chieffi D, Fanelli F, Cho G-S, Schubert J, Blaiotta G, Franz CMAP,
Bania J, Fusco V: Novel insights into the enterotoxigenic
potential and genomic background of Staphylococcus aureus
isolated from raw milk. Food Microbiol 2020, 90:103482.
32. Li L, Renye JA, Feng L, Zeng Q, Tang Y, Huang L, Ren D, Yang P:
Characterization of the indigenous microflora in raw and
pasteurized buffalo milk during storage at refrigeration
www.sciencedirect.com
Microbial community in non-bovine milk Wei et al. 57
temperature by high-throughput sequencing. J Dairy Sci 2016,
99:7016-7024.
33. Silva LF, Sunakozawa TN, Amaral DMF, Casella T, Nogueira MCL,
De Dea Lindner J, Bottari B, Gatti M, Penna ALB: Safety and
technological application of autochthonous Streptococcus
thermophilus cultures in the buffalo Mozzarella cheese. Food
Microbiol 2020, 87:103383.
34. Moossavi S, Azad MB: Origins of human milk microbiota: new
evidence and arising questions. Gut Microbes 2019,
12:1667722.
35. Mosher JJ, Bowman B, Bernberg EL, Shevchenko O, Kan J,
Korlach J, Kaplan LA: Improved performance of the PacBio
SMRT technology for 16S rDNA sequencing. J Microbiol
Methods 2014, 104:59-60.
36. Zhang M, Dang N, Ren D, Zhao F, Lv R, Ma T, Bao Q, Menghe B,
 Liu W: Comparison of bacterial microbiota in raw mare’s milk
and koumiss using PacBio single molecule real-time
sequencing technology. Front Microbiol 2020, 11:2708
This research article applied PacBio single-molecule real-time (SMRT)
sequencing to profile full-length 16S rRNA genes in milk samples, which
enables bacterial taxonomic assignment to the precision of species level.
This is a helpful reference for the accurate assessment of microbial
community in non-bovine milk.
37. Mo L, Yu J, Jin H, Hou Q, Yao C, Ren D, An X, Tsogtgerel T,
Zhang H: Investigating the bacterial microbiota of traditional
fermented dairy products using propidium monoazide with
single-molecule real-time sequencing. J Dairy Sci 2019,
102:3912-3923.
Current Opinion in Food Science 2021, 40:51–57