1709

Journal of Food Protection, Vol. 75, No. 9, 2012, Pages 1709–1714

doi:10.4315/0362-028X.12-101
Copyright G, International Association for Food Protection

Research Note

Molecular Characterization of Antibiotic-Resistant Salmonella

Isolates from Retail Meat from Markets in Northern Vietnam
TRUONG HA THAI1,2 AND RYOJI YAMAGUCHI1*

1Department of Veterinary Pathology, Faculty of Agriculture, University of Miyazaki, Miyazaki 889-2192, Japan; and 2Department of Microbiology–
Infectious Diseases–Pathology, Faculty of Veterinary Medicine, Hanoi University of Agriculture, Trau Quy, Gia Lam, Ha Noi, Vietnam

MS 12-101: Received 28 February 2012/Accepted 28 April 2012

ABSTRACT
A total of 118 Salmonella isolates were detected from 283 retail meat samples (135 pork and 148 chicken meat) purchased at
retail markets in Northern Vietnam. Thirteen serovars, including Infantis, Anatum, Rissen, Reading, Emek, Typhimurium,
Blockley, London, Newport, Derby, Weltevreden, Albany, and Hadar, were determined. Resistance to tetracycline (54.2%),
sulfonamides (52.5%), streptomycin (41.5%), trimethoprim (36.4%), chloramphenicol (35.6%), and ampicillin (33.1%) was
commonly seen in the Salmonella isolates. Fourteen [blaTEM, blaOXA-1, blaPSE-1, aadA1, sul1, tetA, tetB, tetG, cmlA1, floR, dfrA1,
dfrA12, aac(3)-IV, and aphA1-1AB] of 17 resistance genes were detected from the isolates demonstrating resistance. Genes for
plasmid-mediated quinolone resistance, such as qnrA, qnrB, qnrS, qepA, and acc(69)-1b-cr, were not detected in 23 quinolone-
resistant isolates. The substitution TCC to TTC at codon 83 of gyrA was found in the 18 quinolone-resistant isolates. The data
revealed that resistant Salmonella strains were widely distributed in Northern Vietnam via the food chain and that they might
contain multiple genes specifying identical resistance phenotypes. Thus, further studies are necessary to clarify the mechanisms
of antibiotic resistance in Salmonella strains and their spread in the livestock market.

Salmonellosis is one of the most important enteric
diseases, affecting many people worldwide. The global
human health impact of nontyphoidal Salmonella bacteria
is high, with an estimated 93.8 million illnesses annually,
of which an estimated 80.3 million are foodborne and
associated with 155,000 deaths (18). Pork, beef, and
chicken meat have been recognized as significant sources
of human salmonellosis. Although more than 2,500 serovars
of Salmonella enterica have been identified, most human
Salmonella infections are caused by a limited number of
serovars, which may vary from country to country and over
time (13).
The emergence of antibiotic-resistant Salmonella has
become a major public health concern. The use of
antibiotics in any venue, including disease treatment and
growth promotion in domestic livestock, can potentially
lead to widespread dissemination of antibiotic-resistant
bacteria. In recent years, studies of Salmonella isolates
from foodstuffs and animals in Vietnam and other countries
have shown that multidrug resistance of Salmonella is
increasing (25, 27–29).
The increased use of fluoroquinolones has led to
increasing resistance to these antibiotics, with rates of
resistance that vary by both organism and geographic region
(10, 15). The main mechanism of quinolone resistance was
believed to arise from chromosomal mutations in genes
* Author for correspondence. Tel: 81-985-58-7271; Fax: 81-985-58-7271;
E-mail: a0d402u@cc.miyazaki-u.ac.jp.
encoding target enzymes, such as DNA gyrase (gyrA and
gyrB) and/or DNA topoisomerase IV (parC and parE), or
affecting drug accumulation (10, 15, 24). Besides mutations in
chromosomal genes, plasmid-mediated quinolone resistance
(PMQR) has also been reported, including qnr-mediated
inhibition of quinolone binding to DNA, a qepA-encoded
efflux pump, and the aac(69)-1b-cr-mediated inactivation of
fluoroquinolones by an acetyltransferase (21, 22, 24).
The aims of this study were to determinate the
antibiotic resistance in Salmonella sp. isolates from retail
meat from markets in Northern Vietnam and to investigate
the genes coding for antibiotic resistance. In addition, the
nalidixic acid–resistant isolates were further characterized
for quinolone resistance mechanisms.
MATERIALS AND METHODS
Sampling. A total of 283 retail meat samples (135 pork and
148 chicken meat) were collected randomly from the retail markets
in three provinces, including Bac Ninh, Ha Noi, and Ha Tay, in
Northern Vietnam from July 2008 to June 2009. Samples (.100 g)
were placed in sterile plastic sampling bags and chilled in an ice
box during transport to the laboratory at Hanoi University of
Agriculture. All samples were analyzed on the day of arrival.
Salmonella isolation and serotyping. Salmonella organisms
were isolated according to the International Organization for
Standardization standard method ISO 6579 (14), with some
modifications. For preenrichment of Salmonella, 25 g of each
sample was homogenized in a sterile bag with 225 ml of buffered
peptone water and incubated at 37uC for 18 to 24 h. Next, 0.1 ml of
 1710

TABLE 1. Prevalence of antibiotic resistance among Salmonella serovars isolated from retail meat
THAI AND YAMAGUCHI

No. (%) of isolates resistant to drug

Infantis Anatum Rissen Reading Emek Typhimurium London Blockley Newport Others Total
Antibiotica (n ~ 22) (n ~ 21) (n ~ 16) (n ~ 13) (n ~ 9) (n ~ 9) (n ~ 7) (n ~ 7) (n ~ 6) (n ~ 8)b (n ~ 118)

A 9 (40.9) 9 (42.9) 7 (43.8) 1 (7.7) 1 (11.1) 6 (66.7) 1 (14.3) — 3 (50.0) 2 (25.0) 39 (33.1)
Ac — — — — — 4 (44.4) — — — — 4 (3.4)
C 7 (31.8) 8 (38.1) 6 (37.5) 3 (23.1) 4 (44.4) 5 (55.6) 2 (28.6) — 4 (66.7) 3 (37.5) 42 (35.6)
Cf — — — — — — — — — — —
Ci — 1 (4.8) — — — 5 (55.6) — — — 1 (12.5) 7 (5.9)
G 2 (9.1) 4 (19.0) 4 (25.0) — — 5 (55.6) — — 2 (33.3) 1 (12.5) 18 (15.3)
K 10 (45.5) 5 (23.8) 5 (31.3) — — 6 (66.7) — — 4 (66.7) 1 (12.5) 31 (26.3)
Na 4 (18.2) 6 (28.6) 3 (18.8) — — 6 (66.7) — — 3 (50.0) 1 (12.5) 23 (19.5)
Ne 1 (4.5) 3 (14.3) 3 (18.8) — — 6 (66.7) — — 3 (50.0) 1 (12.5) 17 (14.4)
No — — — — — 1 (11.1) — — — — 1 (0.8)
Su 18 (81.8) 7 (33.3) 11 (68.8) 3 (23.1) 5 (55.6) 6 (66.7) 2 (28.6) 1 (14.3) 4 (66.7) 5 (62.5) 62 (52.5)
S 17 (77.3) 7 (33.3) 8 (50.0) 2 (15.4) 1 (11.1) 6 (66.7) 1 (14.3) 1 (14.3) 3 (50.0) 3 (37.5) 49 (41.5)
Te 14 (63.6) 12 (57.1) 10 (62.5) 6 (46.2) 2 (22.2) 6 (66.7) 3 (42.9) 2 (28.6) 6 (100) 3 (37.5) 64 (54.2)
Tp 12 (54.5) 6 (28.6) 5 (31.3) 2 (15.4) 4 (44.4) 5 (55.6) 1 (14.3) — 5 (83.3) 3 (37.5) 43 (36.4)
a
A, ampicillin; Ac, amoxicillin–clavulanic acid; C, chloramphenicol; Cf, ceftazidime; Ci, ciprofloxacin; G, gentamicin; K, kanamycin; Na, nalidixic acid; Ne, neomycin; No, norfloxacin; Su,
sulfonamides; S, streptomycin; Te, tetracycline; Tp, trimethoprim; —, not found.
b
Other serovars included Derby (n ~ 4), Weltevreden (n ~ 2), Albany (n ~ 1), and Hadar (n ~ 1).
J. Food Prot., Vol. 75, No. 9
J. Food Prot., Vol. 75, No. 9 ANTIBIOTIC RESISTANCE OF SALMONELLA FROM RETAIL MEAT IN NORTH VIETNAM 1711

TABLE 2. Distribution and prevalence of resistance genes among the Salmonella serovars
No. (%) of isolates
Drug (no. of isolates tested) and resistance gene(s) positive for gene(s) Serovar (no. of isolates positive for gene[s])

Ampicillin (39)
blaTEM 36 (92.3) Anatum (9), Derby (1), Emek (1), Infantis (7), London
(1), Newport (3), Reading (1), Rissen (5),
Typhimurium (6)
blaOXA-1 9 (23.1) Anatum (2), Derby (1), Infantis (1), Rissen (2),
Typhimurium (3)
blaPSE-1 2 (5.1) Derby (1), Infantis (1)
blaTEMzblaOXA-1 5 (12.8) Anatum (2), Typhimurium (3)
blaTEMzblaPSE-1 1 (2.6) Derby (1)
No. (%) of isolates with at least one gene 39 (100)
Chloramphenicol (42)
cmlA1 24 (57.1) Anatum (6), Derby (1), Emek (1), Infantis (3), London
(1), Newport (4), Rissen (4), Typhimurium (4)
floR 29 (69.0) Anatum (2), Derby (2), Infantis (6), London (1), Newport
(4), Reading (3), Rissen (6), Typhimurium (4),
Weltevreden (1)
cmlA1zfloR 24 (51.1) Anatum (1), Derby (1), Infantis (2), Newport (3), Rissen
(4), Typhimurium (3)
No. (%) of isolates with at least one gene 39 (92.9)
Gentamycin (18)
aac(3)-IV 17 (94.4) Anatum (4), Derby (1), Infantis (2), Newport (2), Rissen
(4), Typhimurium (4)
Kanamycin (31)
aphA1-1AB 31 (100) Anatum (5), Derby (1), Infantis (10), Newport (4), Rissen
(5), Typhimurium (6)
Streptomycin (49)
aadA1 41 (83.7) Anatum (7), Derby (3), Emek (1), Infantis (13), Newport
(3), Reading (1), Rissen (8), Typhimurium (5)
Sulfonamides (62)
sul1 47 (75.8) Anatum (7), Derby (3), Emek (2), Infantis (14), London
(1), Newport (3), Reading (1), Rissen (9),
Typhimurium (6), Weltevreden (1)
Tetracycline (64)
tetA 47 (73.4) Anatum (7), Derby (1), Emek (1), Infantis (10), London
(2), Newport (5), Reading (5), Rissen (9),
Typhimurium (6), Weltevreden (1)
tetB 1 (1.6) Anatum (1)
tetG 20 (31.3) Anatum (4), Derby (1), Infantis (2), Newport (4),
Reading (1), Rissen (3), Typhimurium (4),
Weltevreden (1)
tetAztetB 1 (1.6) Anatum (1)
tetAztetG 16 (25.0) Anatum (2), Derby (1), Infantis (1), Newport (3),
Reading (1), Rissen (3), Typhimurium (4),
Weltevreden (1)
No. (%) of isolates with at least one gene 52 (81.3)
Trimethoprim (43)
dfrA1 20 (46.5) Anatum (3), Derby (1), Emek (2), Infantis (4), London
(1), Reading (2), Rissen (4), Typhimurium (2),
Weltevreden (1)
dfrA12 20 (46.5) Anatum (2), Derby (1), Infantis (6), Newport (3),
Reading (1), Rissen (3), Typhimurium (4)
dfrA1zdfrA12 7 (16.3) Anatum (1), Newport (1), Reading (1), Rissen (2),
Typhimurium (2)
No. (%) of isolates with at least one gene 37 (86.0)
TABLE 3. Resistance profiles and characteristics of quinolone resistance in Salmonella sp. isolates from retail meats a
1712

Disk diffusionc
Characterization MIC (mg/ml)
Serovar of isolate Sourcea of gyrA of Nab Ci No Resistance phenotyped Resistance genes

Anatum C Wild type $32 S S A C Te Na blaTEM, floR

Anatum P Wild type $32 S S A S Te Tp Na blaTEM, tetA, aadA1
Infantis C Wild type $32 S S A C S Su Te K Na blaTEM, floR, aadA1, sul1, tetA, aphA1-1AB,
Infantis P Wild type $64 S S C S Su Te K Ne Na floR, sul1, aadA1, tetA, aphA1-1AB
Anatum C Wild type $64 S S A C S Su Te K Tp Na blaTEM, cmlA1, aadA1, sul1, tetA, tetG, aphA1-1AB, dfrA1
Infantis P Ser83Phe $128 S S C S Su G Tp Na cmlA1, floR, aadA1, sul1, aac(3)-IV, dfrA12
Typhimurium P Ser83Phe $128 S S A S Su Te K Ne Na blaTEM, aadA1, sul1, tetA, aphA1-1AB
THAI AND YAMAGUCHI

Infantis C Ser83Phe $128 S S A C S Su Te K Tp Na blaTEM, cmlA1, aadA1, sul1, tetA, tetG, aphA1-1AB, dfrA12
Newport P Ser83Phe $128 S S A C S Te K Ne Tp Na blaTEM, cmlA1, floR, aadA1, tetA, aphA1-1AB, dfrA1
Rissen C Ser83Phe $128 S S A S Su TK Ne Tp Na blaTEM, aadA1, sul1, tetA, tetG, aphA1-1AB, dfrA1
Anatum C Ser83Phe $128 S S A C S Su Te G K Ne Na blaTEM, blaOXA-1, cmlA1, floR, aadA1, sul1, tetG, aac(3)-IV,
aphA1-1AB
Anatum C Ser83Phe $128 S S A C S Su Te G K Ne Na blaTEM, blaOXA-1, cmlA1, aadA1, sul1, tetG, aac(3)-IV, aphA1-1AB
Newport C Ser83Phe $256 I S A C S Su Te G K Ne Tp Na blaTEM, cmlA1, floR, aadA1, sul1, tetA, tetG, aac(3)-IV, aphA1-
1AB, dfrA1, dfrA12
Newport C Ser83Phe $256 I S A C S Su Te G K Ne Tp Na blaTEM, cmlA1, floR, aadA1, sul1, tetA, tetG, aac(3)-IV, aphA1-
1AB, dfrA12
Rissen C Ser83Phe $256 S S A C S Su Te G K Ne Tp Na blaTEM, cmlA1, floR, aadA1, sul1, tetA, tetG, aac(3)-IV, aphA1-
1AB, dfrA1, dfrA12
Rissen C Ser83Phe $512 S I A C S Su Te G K Ne Tp Na blaTEM, cmlA1, floR, aadA1, sul1, tetA, tetG, aac(3)-IV, aphA1-
1AB, dfrA1, dfrA12
Anatum C Ser83Phe $512 R S A C S Su Te G K Ne Tp Na Ci blaTEM, cmlA1, aadA1, sul1, tetA, tetB, aac(3)-IV, aphA1-1AB,
dfrA1, dfrA12
Derby C Ser83Phe $512 R S A C S Su Te G K Ne Tp Na Ci blaTEM, cmlA1, floR, aadA1, sul1, tetA, tetG, aac(3)-IV, aphA1-
1AB, dfrA12
Typhimurium C Ser83Phe $512 R I A C S Su Te G K Ne Tp Na Ci blaTEM, cmlA1, aadA1, sul1, tetA, tetG, aac(3)-IV, aphA1-1AB, dfrA12
Typhimurium C Ser83Phe $512 R S A C S Su Te G K Ne Tp Na Ci Ac blaTEM, blaOXA-1, cmlA1, floR, aadA1, sul1, tetA, tetG, aac(3)-IV,
aphA1-1AB, dfrA1, dfrA12
Typhimurium P Ser83Phe $512 R I A C S Su Te G K Ne Tp Na Ci Ac blaTEM, blaOXA-1, floR, sul1, tetA, tetG, aphA1-1AB, dfrA12
Typhimurium P Ser83Phe $512 R I A C S Su Te G K Ne Tp Na Ci Ac blaTEM, cmlA1, floR, aadA1, sul1, tetA, aac(3)-IV, aphA1-1AB,
dfrA12
Typhimurium C Ser83Phe $512 R R A C S Su Te G K Ne Tp Na No blaTEM, blaOXA-1, cmlA1, floR, aadA1, sul1, tetA, tetB, tetG,
Ci Ac aac(3)-IV, aphA1-1AB, dfrA1, dfrA12
a
C, chicken meat; P, pork meat.
b
Na, nalidixic acid.
c
Ci, ciprofloxacin; No, norfloxacin; S, susceptible; I, intermediate; R, resistant.
d
A, ampicillin; C, chloramphenicol; Te, tetracycline; Na, nalidixic acid; S, streptomycin; Tp, trimethoprim; Su, sulfonamides; K, kanamycin; Ne, neomycin; G, gentamicin; Ci, ciprofloxacin; Ac,
amoxicillin–clavulanic acid.
J. Food Prot., Vol. 75, No. 9
J. Food Prot., Vol. 75, No. 9 ANTIBIOTIC RESISTANCE OF SALMONELLA FROM RETAIL MEAT IN NORTH VIETNAM
the preenriched culture in buffered peptone water was added to
10 ml of Rappaport-Vassiliadis soya broth, followed by further
incubation at 41.5uC for 24 h. A loopful of culture broth was then
sampled from the selective enrichment Rappaport-Vassiliadis soya
broth, streaked onto xylose lysine Tergitol 4 agar (Merck), and
incubated at 37uC for 24 h. One or two black presumptive colonies
were selected from each plate and cultured on nutrient agar slants.
The isolates were confirmed to be Salmonella by confirmatory
biochemical tests (fermentation of glucose and sucrose, lack of
fermentation of lactose, hydrogen sulfide production test, citrate
test, lysine decarboxylation, and methyl red and indole tests).
Typical Salmonella isolates were serotyped by slide and microtiter
agglutination for O and H antigens (Difco Laboratories, Detroit,
MI) according to the Kauffmann and White scheme (11) by the
Department of Veterinary Hygiene, National Institute of Veterinary
Research, Vietnam.
Antibiotic susceptibility testing. The antibiotic susceptibil-
ities of isolates were determined according to the guidelines of the
Clinical and Laboratory Standards Institute (5). Disk diffusion
assays were performed on Mueller-Hinton agar with disks
containing 14 different antibiotic agents (Oxoid, Basingstoke,
Hampshire, UK). The following antibiotics were tested: ampicillin,
10 mg; amoxicillin–clavulanic acid, 20/10 mg; ceftazidime, 30 mg;
chloramphenicol, 30 mg; ciprofloxacin, 5 mg; gentamicin, 10 mg;
kanamycin, 30 mg; nalidixic acid, 30 mg; neomycin, 10 mg;
norfloxacin, 10 mg; streptomycin, 10 mg; tetracycline, 30 mg;
trimethoprim, 5 mg; and sulfonamides, 300 mg. The interpretive
category susceptible, intermediate, or resistant was determined
according to Clinical and Laboratory Standards Institute guidelines
(7). An isolate was defined as being multiresistant if it was resistant
to three or more antibiotics. Escherichia coli ATCC 25922 was
used as the control organism.
Detection of resistance genes. DNA templates used for PCR
were prepared by boiling bacterial cultures (23). The following
genes implicated in antibiotic resistance were detected by PCR
amplification: blaPSE-1, blaOXA-1, and blaTEM encoding b-lactam
resistance; aadA1, aadA2, aac(3)-IV, aphA-1AB, and Kn encoding
aminoglycoside resistance; catA1, cmlA1, and floR encoding
chloramphenicol resistance; sul1 encoding sulfonamide resistance;
tetA, tetB, and tetG encoding tetracycline resistance; and dfrA1 and
dfrA12 encoding trimethoprim resistance. In the case of an isolate
showing resistance to quinolone-based antibiotic compounds, the
MIC of nalidixic acid was examined by the broth dilution method
(6). The PMQR genes, such as qnrA, qnrB, qnrS, qepA, aac(69)-
1b-cr, and quinolone resistance determining regions (QRDRs) of
gyrA, were detected for quinolone resistance. PCR amplification
reactions were performed in a 25-ml reaction mixture containing
12.5 ml of 2| GoTaq Green master mix (Promega, Madison, WI),
1 ml (10 ng/ml) of primers, 4 ml of DNA template, and nuclease-free
water. The primer sets and the assay conditions used for
amplification were previously described (4, 8, 9, 12, 16, 20).
Sequencing of gyrA. PCR amplifications of the QRDRs to
detect mutations at codons Gly 81, Ser 83, and Asp 87 in gyrA
were carried out using the primers in a previous report (8). Purified
PCR products were sequenced and compared with the wild-type
sequences (GenBank accession no. X78977.1).
RESULTS AND DISCUSSION
Of the 283 retail meat samples, 118 (41.7%) were
Salmonella positive, including 63 (42.6%) of the 148
chicken samples and 55 (40.7%) of the 135 pork samples.
1713
Thirteen serovars were identified, including Infantis,
Anatum, Rissen, Reading, Emek, Typhimurium, Blockley,
London, Newport, Derby, Weltevreden, Albany, and Hadar
(Table 1). Resistance to tetracycline (54.2%), sulfonamides
(52.5%), streptomycin (41.5%), trimethoprim (36.4%),
chloramphenicol (35.6%), and ampicillin (33.1%) was
commonly seen in the Salmonella isolates (Table 1). These
findings were comparable to those in previous reports in
Vietnam (25, 27) and other Asian countries (3, 28, 29). These
drugs have been commonly used for treatment of Salmonella
infection in both humans and animals in these countries
because replacing these antibiotics with the new generations
of drugs is not easy due to their high cost (26, 28).
Fourteen [blaTEM, blaOXA-1, blaPSE-1, aadA1, sul1, tetA,
tetB, tetG, cmlA1, floR, dfrA1, dfrA12, aac(3)-IV, and aphA1-
1AB] of 17 resistance genes were detected from the resistant
isolates (Table 2). In most cases, resistance genes were detected
where the corresponding individual resistance phenotypes were
observed, suggesting the expression of the genes present. High
levels of prevalence of the blaTEM (92.3%), floR (69%), sul1
(75.8%), and dfrA1 (46.5%) genes were observed in the
antibiotic-resistant isolates, similar to a previous report in
Germany (19). In addition, the tetracycline-resistant isolates
were mediated mainly by tetA (73.4%), consistent with other
reports (1, 19). In this study, the mechanism for kanamycin
resistance in the Salmonella isolates was mainly encoded by the
aphA1-IAB gene. However, this gene was infrequent in the
resistant strains of Salmonella Infantis (23).
Resistance of Salmonella isolates to nalidixic acid
(19.5%) in this study was consistent with the findings of
Van et al. (25). However, this level was lower than that in
previous reports in Vietnam (27) and other Asian countries
(3, 28). The low levels of resistance to drugs of the
fluoroquinolone group (norfloxacin and ciprofloxacin) and
third-generation cephalosporins (ceftazidime) were similar
to those in previous reports in Vietnam (25, 27) and other
countries (3, 29). To date, epidemiological data relating to
PMQR genes and QRDRs in Salmonella have been limited
in Vietnam. In this study, PMQR genes, such as qnrA, qnrB,
qnrS, qepA, and acc(69)-1b-cr, were not detected from 23
quinolone-resistant isolates (Table 3), consistent with a
previous report from South Korea (17). However, PMQR
genes were detected at very low rates from quinolone-
resistant Salmonella isolates in France (4), Japan (2), and the
United States (10). Substitutions in the codons of gyrA, such
as Gly 81, Ser 83, and Asp 87, were frequently observed in
Salmonella isolates when the MICs of nalidixic acid were
$128 ml/ml (8, 17). However, no substitutions at codons Gly
81 and Asp 87 in the QRDR of gyrA were identified, but the
single mutation at codon Ser 83 (Ser83Phe) was detected
from 18 of 23 quinolone-resistant isolates in this study
(Table 3). In addition, all of the quinolone-resistant isolates in
this study showed multidrug resistance and carried multiple
resistance genes. This was also frequently observed in the
Salmonella isolates from foodstuffs in Vietnam (27) and
other Asian countries (26, 28) and may result in failure in
salmonellosis treatment in these countries.
Our results showed a high prevalence of antibiotic
resistance among Salmonella strains isolated from retail
1714 THAI AND YAMAGUCHI
meat in Northern Vietnam and correlation between the
genes and the corresponding individual resistance pheno-
types. In addition, the data describe the chromosomal
mutations in nalidixic acid–resistant isolates that may have
resulted from the use of quinolones in animal husbandry.
Therefore, appropriate management strategies are needed to
prevent resistant bacterial pathogens in the food chain in
Vietnam.
ACKNOWLEDGMENTS
This study was supported by the TRIG project of Hanoi Agricultural
University, Gia Lam, Hanoi, Vietnam. We thank the staffs of the
Department of Veterinary Hygiene, National Institute of Veterinary
Research, Hanoi, Vietnam, for their technical assistance with serovar
identification. The authors thank the students, technicians, butchers, and
other individuals who helped with sampling and processing.
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