Experimental Parasitology 135 (2013) 223226

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Experimental Parasitology
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Research Brief

Molecular characterization of Giardia duodenalis isolates from police

and farm dogs in China
Wei Li a,b, Chengwu Liu c, Yuqiang Yu b, Jianhua Li b, Pengtao Gong b, Mingxin Song a, Lihua Xiao d,,
Xichen Zhang b,
a

College of Veterinary Medicine, Northeast Agricultural University, 59 Mucai Street, Harbin 150030, China
College of Animal Science and Veterinary Medicine, Jilin University, 5333 Xian Road, Changchun 130062, China
Shenyang Police Dog Technical College, 4 Baishan Road, Shenyang 110034, China
d
Division of Foodborne, Waterborne and Environmental Diseases, Centers for Disease Control and Prevention, 1600 Clifton Rd NE, GA 30333, USA
b
c

h i g h l i g h t s
 The police and farm dogs have been infected with Giardia duodenalis in Shenyang.
 Signicantly higher infection rates of farm dogs than police dogs were seen.
 The high occurrence of potentially zoonotic subtype AI-1 in dogs is of public health concern.

a r t i c l e

i n f o

Article history:
Received 15 February 2013
Received in revised form 1 July 2013
Accepted 11 July 2013
Available online 25 July 2013
Keywords:
Giardia duodenalis
Dog
Triosephosphate isomerase
Genotyping
Subtyping

a b s t r a c t
To assess the potential zoonotic transmission of giardiasis from dogs in China, a total of 205 fecal specimens from dogs were screened for Giardia duodenalis using PCR and sequence analysis of the triosephosphate isomerase gene. The prevalence of G. duodenalis in dogs was 13.2% (27/205). The potentially
zoonotic assemblage A and the dog-specic assemblage C was identied in 25 (12.2%) and two (1.0%)
dogs, respectively. All assemblage A isolates belonged to sub-assemblage AI, genotype AI-1. Likewise,
one subtype was found in assemblage C. The high occurrence of potentially zoonotic G. duodenalis subtype AI-1 in dogs that are in close contact with humans is of public health concern.
Published by Elsevier Inc.

1. Introduction
Giardiasis is a major diarrheal disease in humans and domestic
and wild animals worldwide (Ballweber et al., 2010; Feng and Xiao,
2011; Thompson and Smith, 2011). Giardia duodenalis (also known
as Giardia lamblia or Giardia intestinalis) is the species infecting
humans and most mammals and consists of at least eight genetically different assemblages A to H. Among them, assemblages A
and B infect both humans and many species of animals, thus are
considered to be potentially zoonotic, whereas the remaining ones
represent host-specic lineages, with assemblages C and D being
mostly found in dogs, assemblage E in domestic ruminants and
pigs, assemblage F in cats, assemblage G in mice and rats, and
assemblage H in marine mammals (Ballweber et al., 2010; Feng

Corresponding authors. Fax: +1 404 718 4197 (L. Xiao), fax: +86 431 8798 1351
(X. Zhang).
E-mail addresses: lxiao@cdc.gov (L. Xiao), zhangxic@public.cc.jl.cn (X. Zhang).
0014-4894/$ - see front matter Published by Elsevier Inc.
http://dx.doi.org/10.1016/j.exppara.2013.07.009

and Xiao, 2011; Monis et al., 2003; Thompson and Smith, 2011;
Xiao and Fayer, 2008). The contamination of source water by
G. duodenalis from animal reservoir hosts is of increasing public
health concern (Baldursson and Karanis, 2011; Feng et al., 2011;
Karanis et al., 2007; Lobo et al., 2009).
Triosephosphate isomerase (tpi) gene is a commonly used genetic marker for differentiating G. duodenalis isolates at the assemblage and subtype levels (Sprong et al., 2009; Sulaiman et al., 2003;
Xiao and Fayer, 2008). For example, in a recent study in the United
States, three main assemblages A, C, and D were seen in dogs based
on tpi sequence analysis (Scorza et al., 2012). Characterization of
G. duodenalis tpi and other genetic loci has revealed a high
incidence of dog-specic assemblages C and D in kennel and
household canines in Croatia (Beck et al., 2012). Another study
compared three genetic loci (SSU-rDNA, elongation factor 1-alpha,
and tpi) for genotyping and subtyping G. duodenalis isolates and
concluded that tpi was the most appropriate marker for assessing
the zoonotic transmission of G. duodenalis between dogs and
humans (Traub et al., 2004).

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W. Li et al. / Experimental Parasitology 135 (2013) 223226

Thus far, few studies have been conducted to assess the occurrence of G. duodenalis genotypes and subtypes in animals in China.
Dogs, as the most common companion animals, are in close contact
with humans in China. In this study, we have examined the occurrence of G. duodenalis in 205 police and farm dogs and characterized the parasite at both the genotype and subtype levels using
PCR and sequence analysis of the tpi gene.
2. Material and methods
2.1. Ethical considerations
Prior to the collection of canine fecal specimens, permission
was obtained from the farm owners and the police canine managers. Dogs were caged, fed alone in each cage and on the following
day, fecal specimens were collected in plastic bags. The animals
were not harmed in any way during the procedure.
2.2. Specimens
A total of 205 fecal specimens were obtained during October to
December 2011 from police and farm dogs in Shenyang, China.
Among them, 52 specimens were collected in October and November 2011 from ve farms (canine breeding facilities) in suburb and
rural areas of Shenyang, where the dogs were free ranging. A second collection of 153 specimens was done in December 2011 at
a police canine training station in urban Shenyang, where dogs
were kept in cages. Only one specimen per dog was used in the
study. All the dogs sampled had frequent contact with their keepers or trainers, and the farm dogs had access to source water. The
dogs were assigned into two age groups: adult group with animals
older than one year and juvenile group with animals aged between
two months to one year.
2.3. Detection of Giardia infection by PCR
Fecal specimens were washed twice in distilled water, and
genomic DNA was extracted from 0.3 g of washed specimens using
a QIAamp DNA Stool Mini Kit (QIAGEN, Hilden, Germany) and
manufacturer-recommended procedures. G. duodenalis in specimens was detected by nested PCR amplication of a 532-bp fragment of the tpi gene as described (Sulaiman et al., 2003). Each
specimen was analyzed twice using 2 ll of the DNA extract per
PCR. Non-acetylated bovine serum albumin (TaKaRa, Japan) at a
concentration of 400 ng/ll was used in primary PCR to neutralize
residual PCR inhibitors in the extracted DNA. PCR products were
visualized by electrophoresis in 1.5% agarose containing ethidium
bromide.
2.4. Genotyping and subtyping of G. duodenalis
The secondary PCR products of the anticipated size were sent to
the Sangon Company (Shanghai, China) for DNA sequencing at both
directions. The nucleotide sequences obtained were edited using
Chromas Pro 1.33 (Technelysium Pty Ltd, Helensvale, Queensland,
Australia) and aligned with reference sequences using the ClustalX
1.81 package (ftp://ftp-igbmc.u-strasbg.fr/pub/ClustalX/) to identify Giardia species, genotypes, and subtypes.
2.5. Statistical analysis
Statistical analysis was conducted using SPSS version 17.0 (SPSS
Inc., Chicago, IL, USA). Infection rates with G. duodenalis were compared using Chi-square analysis at a signicance of p < 0.05.

3. Results
3.1. G. duodenalis occurrence
Overall, 27 of 205 specimens [13.2%; 95% condence interval
(CI): 0.0820.181] produced the expected tpi PCR product, including 15 (9.8%; 95% CI: 0.0480.148) from 153 police canines and 12
(23.1%; 95% CI: 0.1000.361) from 52 farm dogs. The difference in
infection rates between farm dogs (23.1%) and police canines
(9.8%) was signicant (p < 0.05). A signicantly higher infection
rate was seen in adult dogs (26 of 103 or 25.2%; 95% CI: 0.155
0.349) than in juvenile dogs (1 of 102 or 1.0%; 95% CI: 0.009
0.029) (p < 0.01). In the police canine training station, 15 of 70
(21.4%; 95% CI: 0.1060.323) adult dogs examined were positive,
whereas none of the 83 juvenile dogs were positive. On farms, 11
of 33 (33.3%; 95% CI: 0.1360.530) adult dogs and one of 19
(5.3%; 95% CI: 0.0510.156) juvenile dogs examined were positive (p < 0.05). The overall infection rate in female dogs (18/109
or 16.5%; 95% CI: 0.0890.241) was slightly higher than males (9/
96 or 9.4%; 95% CI: 0.0330.155) (p > 0.05). On farms, male dogs
(9/35 or 25.7%; 95% CI: 0.0890.425) had an infection rate higher
than females (3/17 or 17.6%; 95% CI: 0.0230.376) (p > 0.05). In
the police canine station, 15 of 92 (16.3%; 95% CI: 0.0810.246) female dogs were positive, whereas none of the 61 male dogs were
positive.
3.2. G. duodenalis genotypes and subtypes
Sequence analysis of tpi PCR products identied G. duodenalis
assemblages A and C. The former was detected in 25 (12.2%; 95%
CI: 0.0740.170) specimens, of which 10 were from farm dogs
and 15 from police canines. Within assemblage A, one subtype
was identied: sub-assemblage AI, genotype AI-1, previously
found in humans, cattle, water buffaloes, cats, pigs, dogs, and sheep
(Feng and Xiao, 2011). Subtype AI-1 was found in seven male adult
dogs, one male juvenile, and two female adult dog on farms and 15
female adult police dogs (Table 1). In contrast, assemblage C was
only seen in two (1.0%; 95% CI: 0.0040.023) specimens from
farm dogs (Table 1). The two assemblage C specimens had tpi
sequences that were identical to a sequence previously identied
in a dog in Atlanta, USA (GenBank accession No. AY228643)
(Sulaiman et al., 2003) (Table 2). The subtype identied was named
as CI-1 in this study (Tables 1 and 2).
4. Discussion
Infection rates of G. duodenalis in dogs were reportedly high in
some countries: 36.8% in Brazil, 26.6% in Italy, 23.4% in Japan,
22.7% in Belgium, and 21.0% in United Kingdom (Claerebout
et al., 2009; Itoh et al., 2011; Paoletti et al., 2008; Upjohn et al.,
2010; Volotao et al., 2007). Lower infection rates were reported
in some other countries such as the Netherlands (15.2%), Italy
(15.0%), Peru (14.5%), China (11.0%), Australia (9.4%), Nicaragua
(8.0%), Thailand (7.9%), Finland (5.3%), and Poland (2.0%) (Berrilli
et al., 2004; Cooper et al., 2010; Inpankaew et al., 2007; Lebbad
et al., 2008; Li et al., 2012; Overgaauw et al., 2009; Palmer et al.,
2008; Rimhanen-Finne et al., 2007; Solarczyk and Majewska,
2010). The data obtained in this study clearly showed higher infection rates of G. duodenalis in free range dogs on farms than caged
dogs in a police canine training station. This may result from the
more frequent contact among farm dogs. Most earlier studies have
shown lower infection rates in adult animals than in juveniles
(Ballweber et al., 2010; Feng and Xiao, 2011; Xiao and Fayer,
2008). Thus, one recent study from China reported that the infection rate of G. duodenalis was signicantly higher in young pet dogs

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W. Li et al. / Experimental Parasitology 135 (2013) 223226

Table 1
Giardia duodenalis genotypes and subtypes based on the tpi gene characterization in police and farm dogs in Shenyang, China.
Locations

No. of specimens

Farms
Police canine training station
Total
a

52
153
205

Assemblage A

Assemblage C
a

# Positive

Subtype (# positive and gender and age )

# Positive

Subtype (# positive and gender and agea)

10
15
25

AI-1 (7 MA, 1 MJ, and 2 FA)

AI-1 (15 FA)
AI-1 (7 MA, 1 MJ, and 17 FA)

2
0
2

CI-1 (1 MA and 1 FA)

CI-1 (1 MA and 1 FA)

MA: male adult, MJ: male juvenile, and FA: female adult.

Table 2
Nucleotide sequence differences in the partial tpi gene among subtypes of G. duodenalis assemblage C.

Sequences

Locations

References

AY228641
AY228642
AY228643
AY228644
HQ696782
AF069563
EU781004
EU781005
EU781006
EU781007
CI-1

USA
USA
USA
USA
USA
Australia
Sweden
Sweden
Sweden
Sweden
China

Sulaiman et al. (2003)

Sulaiman et al. (2003)
Sulaiman et al. (2003)
Sulaiman et al. (2003)
Wang et al. (2012)
Monis et al. (1999)
Lebbad et al. (2010)
Lebbad et al. (2010)
Lebbad et al. (2010)
Lebbad et al. (2010)
This study

Nucleotide at positiona
36

72

82

126

207

252

284

305

315

330

T
T
T
T
T
T
T
T
C
C
T

G
G
G
T
G
G
G
T
G
G
G

G
G
G
G
G
G
G
G
G
G
G

C
C
C
C
C
C
C
C
C
C
C

G
G
G
G
G
G
G
G
G
G
G

T
C
C
C
T
T
T
C
T
T
C

G
G
G
G
G
G
G
G
G
G
G

C
C
T
T
C
C
C
T
C
C
T

A
A
C
C
A
A
A
C
A
A
C

G
G
G
G
G
G
G
G
A
G
G

Nucleotide position numbers according to AY228642, with the beginning of the rst nucleotide as position No. 1.

less than six months (25.58%) than in dogs of six months to three
years (7.37%) and adults over three years (7.04%) (Li et al., 2012).
The reason for the difference between the current and earlier studies is not clear.
In most studies on G. duodenalis in dogs, assemblages A, C, and D
were the dominant assemblages (see the review by Feng and Xiao,
2011 on earlier data). The dog-specic assemblages C and D were
far more prevalent than the zoonotic assemblage A in most areas
(Beck et al., 2012; Berrilli et al., 2012; Feng and Xiao, 2011; Itoh
et al., 2011; Liang et al., 2012; McDowall et al., 2011). In contrast,
in this study, assemblage A was more commonly found in dogs
than assemblage C. Assemblage D was not found in any of the dogs,
but this could be due to the non-amplication of the tpi gene of
this groups of parasites by the PCR primers used (Lebbad et al.,
2010). In a few recent studies, assemblage A was also the dominant
genotype of G. duodenalis in dogs (Schurer et al., 2012; Volotao
et al., 2011). A recent study on G. duodenalis in pet dogs in China
had also identied both assemblages A (in 5 animals) and D (in
18 animals) (Li et al., 2012).
Subtyping of assemblage A at various loci has identied at least
three broad sub-assemblages: AI, AII, and AIII. Within each of these
sub-assemblages, various genotypes have been identied, with
sub-assemblage AI (genotype A1) detected in a range of hosts
including humans, cattle, water buffaloes, sheep, dogs, cats, pigs,
and other animals. Various genotypes of AII circulate mainly in humans and AIII mostly wild ruminants (Caccio et al., 2008; Feng and
Xiao, 2011). In China, both AI-1 and AII have been identied in humans (Wang et al., 2011). In this study, most of the assemblage A
infections in dogs was caused by subtype AI-1. Although this subtype is more commonly seen in animals than in humans (Feng and
Xiao, 2011), due to the close contact between dogs and humans in
China, the frequent nding of subtype AI-1 is of potential public
health signicance. Nevertheless, the observations at the tpi locus
need to be substantiated using other genetic loci.
Acknowledgments
This work was supported by Major Program of Preventive and
Control for National Severe Infective Diseases (No. 2008ZX10004-

001) and funds from Northeast Agricultural University (No.

2012RCA01). The ndings and conclusions in this report are those
of the authors and do not necessarily represent the views of the
Centers for Disease Control and Prevention.
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