C H A P T E R

22
Japanese Quail as a Laboratory
Animal Model
Janet Baer, DVMa, Rusty Lansford, PhDb and
Kimberly Cheng, PhDc
a
California Institute of Technology, Pasadena, CA, USA bChildren’s Hospital Los Angeles, Keck
School of Medicine at USC, Saban Research Institute, Department of Radiology, Los Angeles, CA,
USA cThe University of British Columbia, Faculty of Land and Food Systems, Avian Research
Centre, Vancouver, BC, Canada

O U T L I N E

  I. Introduction 1087 IV. Breeding and Rearing Quail 1097

A. Ecology and Taxonomy 1087 A. Reproduction 1097
B. Use in Research 1088 B. Egg Storage and Incubation 1098
C. Hatching 1099
 II. Biology 1089
D. Rearing Chicks 1100
A. Morphology and Physiology 1089
B. Behavior 1091  V. Diseases/Welfare Concerns 1100
C. Genetics 1093 A. Noninfectious 1100
B. Infections 1101
III. Laboratory Management and Husbandry 1094
A. Sources of Quail 1094 References 1104
B. Housing Systems 1094
C. Food and Water 1095
D. Common Procedures 1096

I. INTRODUCTION modest size of breeding adults, ease of breeding in labo-

Japanese quail (Coturnix japonica) are used as a lab-
oratory animal model for multiple areas of scientific
inquiry including, but not limited to developmental
biology, endocrinology, aging, immunology, behavior
studies, and a variety of human genetic disorders. The
quail embryo is an amniote with early developmental
patterns remarkably similar to those of humans; as such
they present significant experimental advantages for the
study of amniotes, e.g., rapid reproductive maturation,
ratory animal facilities, resilience to research manipula-
tions, availability of transgenic lines, a fully sequenced
genome, and tools for molecular manipulations.
A.  Ecology and Taxonomy
Japanese quail are terrestrial birds that generally live
for 2–3 years in the wild (Ottinger, 2001). Their natu-
ral habitat consists of ever-dwindling grasslands, crop-
lands, riversides, mountain slopes near water, and grass

Laboratory Animal Medicine, Third Edition

DOI: http://dx.doi.org/10.1016/B978-0-12-409527-4.00022-5 1087 2015 Elsevier Inc. All rights reserved.
© 2012
1088 22.  Japanese Quail as a Laboratory Animal Model
steppes in eastern Asia, including northern Mongolia,
eastern Russia, northeastern China, Japan, South Korea,
and North Korea (del Hoyo et  al., 1994; Bump, 1971).
They migrate seasonally; however, the migration pat-
tern is complicated and not well understood. Some
populations in Japan are year-round denizens, but most
Japanese quail migrate south to winter in southern
China, Vietnam, Laos, Myanmar, Cambodia, Bhutan, and
northeastern India (del Hoyo et al., 1994; Pappas, 2002).
Once thought to be common in China (del Hoyo et  al.,
1994), decreased quail populations seem to be occurring
in Laos (Duckworth, 2009), Japan (Okuyama, 2004), and
possibly throughout their habitat (del Hoyo et al., 1994;
Duckworth, 2009). Endemic populations of Japanese
quail are listed as ‘Near Threatened’ on the IUCN Red
List (IUCN, 2013a) because they appear to have under-
gone an 80% population decline between 1973 and 2002,
potentially owing to hunting (Okuyama, 2004), shifts in
agriculture (Duckworth, 2009; IUCN, 2013b), contamina-
tion of the wild gene pool by escaped or released farm
quail, and climate change. Research is urgently needed
to establish population numbers and trends, as well as to
determine the threats to naturally occurring wild popu-
lations of Japanese quail. Attempts to introduce them
into the mainland of the United States as a gamebird, in
a previous era when such transplants were considered
a good idea, consistently failed due to a lack of under-
standing of their migratory tendencies (Standford, 1957).
In contrast, introductions to the Hawaiian Islands were
successful (Peterson, 1961).
Japanese quail belong to the order Galliformes, family
Phasianidae, genus Coturnix, and species japonica. They
are classified as Old World quail and are closely related
to the European common quail, Coturnix coturnix. The
two species are not thought to interbreed in native loca-
tions where they co-exist, so they are considered to be
in an intermediate stage of speciation (Johnsgard, 1988;
Pappas, 2002; Howard and Moore, 1984; Kano, 2006;
Crawford, 1990; Union, 1983). However in captivity,
C. japonica and C. coturnix will interbreed and produce
fertile hybrids (Johnsgard, 1988).
B.  Use in Research
1.  Quail as a Model for Embryogenesis Studies
The understanding of myogenesis, vasculogenesis,
angiogenesis, skeletogenesis, virology, immunology,
endocrinology, and teratology has progressed signifi-
cantly as a result of studies in avian embryos (Mizutani,
2002). In Japanese quail, embryogenesis progresses faster
than in the chicken, yet closely mirrors that of the chicken.
On average, incubation times for the quail and chicken
are 16 days and 21 days, respectively. After Hamburger
and Hamilton (Hamburger and Hamilton, 1951) estab-
lished the prototype avian embryo staging system in
LABORATORY ANIMAL MEDICINE
the chicken, outlining the various developmental events
that occur during each stage, Japanese quail embryos
were similarly staged and registered (Padgett and Ivey,
1959, 1960; Zacchei, 1961; Ainsworth et  al., 2010). More
recently, developmental stages of the quail have been
further delineated using modern imaging technologies
(Ruffins et  al., 2007). The accelerated ontogeny of quail
embryos at mid- to late stages of gestation results in loss
of precise registration with the chicken.
Developmental biologists have long used the differ-
ences between chicken and quail embryologic develop-
ment to transplant tissue from one of these into the other
(Le Douarin and Barq, 1969; Le Douarin and Kalcheim,
1999). Quail interphase nuclei have nucleoli with compact
heterochromatin that stains intensely with Schiff’s reagent,
while chicken nucleoli are not stained (Le Douarin, 1973);
thus permitting quail cells to be distinguished from
chicken cells in chimeric embryos. The chick–quail chi-
mera system has been effectively used for a myriad of
cell lineage analyses (Le Douarin and Kalcheim, 1999).
Similarly, quail cells and tissues have been reciprocally
transplanted into other avian species, including ducks to
generate quail–duck chimeras (i.e., qucks), and for heter-
ochrony studies (Lwigale and Schneider, 2008).
The quail embryo can be imaged for minutes to days
in ovo or in vitro using any number of imaging modalities,
e.g., light microscopy, fluorescent microscopy, magnetic
resonance imaging, and computer-assisted tomography.
For light and fluorescent microscopy, the quail embryo
is visible through a windowed egg for continuous imag-
ing over several days (Kulesa et  al., 2000; Bower et  al.,
2011). The quail embryo itself is also physically acces-
sible, which permits cell and tissue transplantations and
genetic manipulations. Differences between the preferred
amniote model for molecular studies (mice), versus the
best model for live imaging (avians), motivated the con-
struction of transgenic, fluorescent protein-expressing
Japanese quail as an experimental system using lentiviral
vectors. Transgenic quail lines expressing fluorescently
labeled cells in a ubiquitous or tissue-specific manner per-
mit amniote embryogenesis to be dynamically recorded
with subcellular resolution (Seidl et  al., 2013; Sato et  al.,
2010; Sato and Lansford, 2013; Scott and Lois, 2005).
2.  Quail as a Model for Health and Disease
Japanese quail are emerging as an animal model to
study birth defects and diseases that affect human health
(Cheng and Kimura, 1990; Mizutani, 2002; Tsudzuki,
2008), and a number of quail lines currently exist that
recapitulate human hereditary diseases, malformations,
and abnormalities. Quail have been used in studies
addressing senescence in immunology, endocrinology,
developmental (Dickman et  al., 2004) and reproductive
biology (Ottinger et al., 2004); hypercholesterolemia char-
acterized by the development of vascular lesions and
 II. Biology
xanthomatosis (Hoekstra et  al., 1998; Murakami et  al.,
2010; Yuan et al., 1997); glycogen storage disease (Matsui
et  al., 1983); and myotonic dystrophy and acid maltase
deficiency, known as Pompe’s disease (Kikuchi et  al.,
1998; Mizutani, 2002). A neurofilament-deficient mutant
of the Japanese quail, named ‘Quiver,’ exhibits wide-
spread tremors (Hasegawa et  al., 1996); in this strain,
neurofilament protein is lacking from neuronal axons
and cell bodies. The noradrenaline and 5-hydroxytryp-
tamine content in the neostriatum of the Quiver’s brain
differs from that in the normal quail, although disap-
pearance of the three neurofilament proteins is observed
in all areas of the Quiver’s brain.
Japanese quail follow a predictable aging process with
evidence of declining function in reproductive, metabolic,
and sensory systems (Holmes and Ottinger, 2003), com-
parable to senescence in mammals. The use of Japanese
quail to study aging provides two distinct advantages
over mammalian models. First, hypothalamic systems
in quail exhibit neuroplasticity; for example, aging
males respond to testosterone replacement therapy with
full recovery of sexual behavior (Ottinger et  al., 2004).
Behavioral recovery is accompanied by the restoration of
specific hypothalamic neuropeptide systems, which reg-
ulate both sexual behavior and gonadotropin-releasing
hormone, thereby allowing identification of these neural
systems. The male cloacal (proctodeal) gland, which is
similar to the prostate gland in mammals, is androgen
responsive. The second advantage to the quail model
is that they exhibit a dynamic bone physiology. This is
particularly true for females because their hollow bones
serve as a reservoir for minerals used in egg production.
Thus, aging female quail develop bone fragility and serve
as a validated model for hormone effects on osteoporosis
and the role of vitamin D (Takahashi et al., 1983). Rescue
of these systems via hormone replacement provides a
tool for studying systems affected by aging and the res-
toration of their functions. Japanese quail have also been
a model system for studying the genetics and neuro-
endocrine control of reproductive and social behavior
(Ball and Balthazart, 2010; Nol et al., 1996), including use
as an animal model for testing the effects of endocrine-
disrupting chemicals (Tattersfield et al., 1997; Hutchinson
et al., 2000). Genetically unique strains of Japanese quail
have been used to study the visual system; developmen-
tal pathology of the retina and optic nerve (Takatsuji
et al., 1984), spontaneous glaucoma (Takatsuji et al., 1986,
Weidner et al., 1995), cataracts (Takatsuji et al., 1985), and
retinal degeneration (Zak et  al., 2010) are some of the
topics that have been investigated.
Japanese quail are susceptible to pathogenic strains
of bacteria such Salmonella pullorum, Escherichia coli
(Woodard et  al., 1973), and Mycobacterium avium (Tell
et  al., 2003), and have served as an animal model for
understanding these pathogens in poultry. Recently
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1089
Rundfeldt et  al. (2013) employed quail to assess the
efficacy of nanoparticulate itraconazole aerosolization
against Aspergillus fumigatus.
II. BIOLOGY
A.  Morphology and Physiology
Japanese quail measure ~17 cm in length, and males
are slightly smaller than females. Whereas wild females
weigh ~100 g and males weigh ~90 g, domesticated adult
females weigh ~140 g and males weigh ~130 g. Some
Japanese quail bred for meat production can weigh
~600 g. The birds have dark-brown feathers from the
top of the head down to the nape of the neck with a
tan streak over the eyes. Adults are sexually dimorphic,
with females having light tan feathers and black speck-
ling on their chest and throat and males characterized
by a rusty brown throat and breast feathers (Fig. 22.1).
Various plumage colors have been prized and selec-
tively bred for over the past five centuries, e.g., rusty
brown, Manchurian Gold, English Black, Barred, White,
and A&M Giant White (Wetherbee, 1961; Roberts, 1999).
Many plumage color mutants have also been developed
and maintained for genetic studies (Cheng and Kimura,
1990) (Fig. 22.2).
The relatively small size and rapid growth rate of
Japanese quail simplifies housing and production in lab-
oratory animal facilities. Body weight at hatching ranges
from 6 to 8 g with growth to 160–250 g by 5–6 weeks of
age, depending on the sex and strain (Gerken and Mills,
1993) (Fig. 22.3). By 4–6 weeks post-hatch, males and
females are sexually dimorphic and can be differentiated
based on plumage coloration. Quail can be sexed within
one day of hatching by cloacal examination; however,
accuracy in performing this technique may take years to
develop (Homma et  al., 1966). Life expectancy in com-
mercial and laboratory settings ranges from 1.5 to 2.5
years, but individual birds have been reported to live
up to 8 years (Cheng et al., 2010).
The cloacal gland of adult male Japanese quail secretes
a white foamy material (Adkins-Regan, 1999; Cheng
et  al., 1989a,b); increased gland size correlates with an
increased photoperiod (Sachs, 1969). Foam deposited
during natural copulation can be retained by the female
for more than 12 h and contributes to prolonged sperm
motility (Biswas et  al., 2010). Cloacal gland size can be
used as an external indicator of testicular function in
male birds (Mohan et  al., 2002), while gland regression
and decreased body weight has been associated with
castration. Observation of foam following gentle manual
pressure of the cloacal gland is also useful for differenti-
ating males from females in strains that have a plumage
color other than wildtype. In addition, foam production
1090 22.  Japanese Quail as a Laboratory Animal Model

FIGURE 22.1  Wildtype plumage of a male (A) and female (B) adult Japanese quail (Coturnix japonica).

FIGURE 22.2  Japanese quail plumage color mutants developed for genetic studies: albino (A), yellow (B), white breasted (C), and silver (D).

has been related to the presence of bacteria in the cloacal

150 gland. Treatment with fluoroquinolones has been dem-
onstrated to result in decreased bacterial counts in the
foam, decreased foam production, and decreased fertil-
ity (Mohan et al., 2004).
Japanese quail have color vision, which appears to be
Body weight (g)

100 Female
more crucial than pattern or form for visual discrimi-
Male nation (Fidura, 1969). Similar to other ground-foraging
birds, quail exhibit a lower field myopia that permits
50 them to simultaneously focus on the ground while they
forage and monitor the horizon and sky overhead for
predators (Hodos and Erichsen, 1990). They exhibit
greatest sensitivity in the auditory range between 1 and
0
4 kHz (Niemiec et  al., 1994). Young chicks (3–5 h post-
hatch) show auditory discrimination learning, respond-
0 2 4 6 8
ing at greater frequency to a familiar sound than a novel
Age (wk)
sound (Evans and Cosens, 1977). Limited information
FIGURE 22.3  Growth curve for male and female domestic Japanese is available regarding the senses of taste and smell in
quail from hatching to 8 weeks of age. Data from Aggrey (2003). Japanese quail (Mills et al., 1997).

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 II. Biology 1091
TABLE 22.1 Standard Biological Data for Domestic Japanese TABLE 22.2 Hematological1 and Clinical Chemistry Values2
Quail for Adult Domestic Japanese Quail
BODY WEIGHT (G) Laying
Adult male females
1 day 6–8
Erythrocytes (106/mm3) 4.14 ± 0.07 3.81 ± 0.14
Adult male 100–130
Packed cell volume (%) 53.1 ± 0.8 46.9 ± 1.3
Adult female 120–160
Haemoglobin (g/100 ml) 15.8 ± 0.2 14.3 ± 0.5
ORGAN WEIGHT (% BODY WEIGHT)
3
Mean corpuscular volume (µm ) 127.0 ± 2.0 124.0 ± 2.0
Liver 1.95
Mean corpuscular haemoglobin (ng) 38.5 ± 0.1 37.7 ± 0.7
Heart 0.91
Mean corpuscular haemoglobin 29.6 ± 0.3 30.4 ± 0.4
Kidney 0.73 concentration (%)

Testes 2.88 Reticulocytes (%) 7.0 ± 0.5 6.1 ± 0.4

3 3
Body temperature ( C) o
40.7–43.2 Thrombocytes (10 /mm ) 117.0 ± 9.0 132.0 ± 17.0

PERFORMANCE AND LONGEVITY Total leucocytes (103/mm3) 19.7 ± 0.7 23.1 ± 1.0
  Heterophils (%) 20.8 ± 1.9 21.8 ± 1.8
Egg weight (g) 9–10
  Eosinophils (%) 2.5 ± 0.04 4.3 ± 1.5
Egg number/100 bird days 80–90
  Basophils (%) 0.4 ± 0.1 0.2 ± 0.1
Age at sexual maturity (d) 38–42
  Lymphocytes (%) 73.6 ± 2.1 71.6 ± 1.
Life span (months) 24–26
  Monocytes (%) 2.7 ± 0.3 2.1 ± 0.3
BLOOD PRESSURE (MMHG)
Glucose (mmol/l)* 17.3 ± 0.6 14.4 ± 0.6
Systolic
Uric acid (mmol/l)* 324 ± 21.5 320 ± 13.9
Adult male 151.8 ± 4.61 Total cholesterol (mmol/l) 6.7 ± 0.13 7.9 ± 0.26
Adult female 156.1 ± 4.7 1
Triglyceride (mmol/l)* 3.0 ± 0.2 23.5 ± 2.2
Diastolic Bilirubin (µmol/l) 20.4 ± 1.00 8.9 ± 0.7
ASAT (U/l) 422 ± 9.5 402 ± 13
Adult male 158.1 ± 4.71
ALAT (U/l) 9.6 ± 0.6 6.5 ± 1.1
Adult female 146.9 ± 4.21
γ-GT (U/l) 1.7 ± 0.2 1.9 ± 0.1
HEART RATE (BEATS/MIN)
Cholinesterase (kU/l) 4.8 ± 0.1 2.8 ± 0.1
Adult male 530.7 ± 17.71
Creatinine (µmol/l)* 4.0 ± 0.4 4.5 ± 0.3
Adult female 489.5 ± 17.11
Protein (g/l) 25.0 ± 1.0 33.6 ± 4.6
ADULT FEED CONSUMPTION (G/DAY) 17.5–19.8 Albumin (g/l) 13.3 ± 0.2 15.3 ± 0.2
Modified from Cheng et al. (2010).
Phosphate (mmol/l) 1.2 ± 0.3 1.9 ± 0.4
1
Mean ± SE.
Calcium (mmol/l) 2.3 ± 0.1 4.0 ± 1.3
Magnesium (mmol/l) 1.0 ± 0.04 1.2 ± 0.1

Standard biological data for Japanese quail are Iron (µmol/l) 12.5 ± 0.4 21.0 ± 2.8
presented in Table 22.1, while hematological and clini- Values given are mean ± SE.
cal chemistry values are given in Table 22.2. Sex related *
Significant daily patterns in concentration (Herichová et al., 2004).
1
Data from Nirmalan and Robinson, 1971.
differences in serum chemistry reference values in 2
Data from Faqi et al., 1997;Scholtz et al., 2009b.
adult Japanese quail have been reported (Scholtz et  al.,
2009a).

in a simulated natural environment found all but two

B. Behavior of the courtship displays and behavioral components
Two studies (Nichols et al., 1991,1992) that compared reported in the literature were exhibited by both groups.
the behavior of feral (considered wild because they were The differences in reproductive behavior between feral
feral since the 1920s) and domesticated Japanese quail and domestic Japanese quail are shown in Table 22.3.

LABORATORY ANIMAL MEDICINE

1092 22.  Japanese Quail as a Laboratory Animal Model

TABLE 22.3 Behavioral Differences between Feral and with the bill, scratching with the legs, tossing dust into
Domestic Japanese Quail in Outdoor Aviary with Simulated the air with the wings while moving the body under
Natural Environment
the dust shower, accompanied by active feather ruffling
Behavior Feral* Domestic and shaking (Mills et  al., 1997). Dust bathing behavior
occurs more frequently in the late afternoon compared
Male crowing Less frequently; varied their More frequently
crowing frequency during with no temporal
to the early morning or early afternoon (Abdelfattah
the female’s laying cycle variation et  al., 2012). In another report, the impact of four types
of environmental enrichment (foraging opportunities,
Courtship More Less
displays structural complexity, sensory stimulation/novelty, and
social housing) were assessed (Miller and Mench, 2005).
Copulations Less frequent More frequent
Use of foraging opportunities, structural complexity and
Male More; toward both males Less dust bathing was observed in 29, 26, and 16% of activ-
aggression and females ity budget scans, respectively, which supports a posi-
Female Less More; toward males tive response to this type of enrichment. Social housing
aggression decreased the use of environmental enrichment and
Pair bond Paired for the whole Shorter; frequently dust baths. Female Japanese quail co-housed with males
duration breeding season switched mates in floor pens used environmental enrichment, foraged
Female laying 1 clutch of eggs for the Up to three clutches more, and were more active than the males.
season for the season Exposure to mild, unpredictable stressors during
Data from Nichols (1991). routine husbandry may impact the morphological and
*
Captured from the feral population on Hawaii. Japanese quail were released on Hawaii behavioral development of quail offspring. Chicks
in the late 1920s. produced by laying hens exposed to mild daily stress-
ors showed altered growth and behavioral responses.
Potential stressors include sudden noise, sudden appear-
The studies indicated that domestication was not ance of bright colored objects, or shaking the cages
‘degenerative,’ but rather behavioral components are (Guibert et al., 2011).
not expressed under conditions that lack the appropri- The sexual behavior of captive Japanese quail ranges
ate stimuli. from polygamy to monogamy (Kovach, 1975; McKinney
Because quail housed in a semi-natural environment et  al., 1983; Orcutt and Orcutt 1976). Females housed
spent approximately 8% of their activity budget on for- continuously with the same male had significantly
aging behavior, including pecking and scratching, it is higher egg hatching rates than females paired with a
suggested that creating the opportunity for birds to per- male every third day. When paired with a male every
form this behavior will improve animal welfare (Schmid 3 days, females paired with the same male had fewer
and Wechsler, 1997). A solid floor environment provided hatched eggs than females paired with a different male.
with a suitable substrate, e.g., wood chips, encourages Males were more receptive to females introduced into
foraging, and opportunities to forage can be created the male’s cage than vice versa (Sullivan et  al., 1992). A
through scattering small seeds or grains in the bedding detailed review of social behavior, including courtship
substrate or through providing other food items in the and vocalization types, rates, and patterns, has been
environment. In contrast, the same study found the birds published (Cheng et al., 2010).
spent little time on elevated structures; hence the pro- Japanese quail show dominance through the devel-
vision of perches is likely of little value. Also of inter- opment of a pecking order (Gerken and Mills, 1993);
est is that quail in semi-natural aviaries stayed under increased stocking density, and changes in group com-
cover for a significant percentage of time (Schmid and position lead to more aggression and pecking-related
Wechsler, 1997). Use of nest boxes with a small entrance injuries. In addition, adult females may outweigh males
but no other openings has been reported (Buchwalder and will direct aggressive behavior toward smaller
and Wechsler, 1997). In outdoor aviaries, this type of males. Maintenance of stable social groups over time
nest box should be kept in the shade, especially in hot improves animal welfare through reducing aggression
climates, as elevated nest box temperatures can be lethal associated with mixing social groups. Multi-male groups
to incubating females. are associated with increased injuries due to aggres-
Captive Japanese quail engage in several dust bathing sive pecking between males (Wechsler and Schmid,
sessions daily when provided with a suitable material 1998). Provision of visual barriers, reducing the age at
such as sand or cat litter (Schein and Statkiewicz, 1983). introduction, and changes in the male: female stocking
In the absence of such material, birds may exhibit ‘vac- density did not reduce male to male induced injuries.
uum dust bathing,’ during which behavioral components Importantly, reducing the light intensity decreased but
of dust bathing are expressed, e.g., raking movements did not eliminate pecking-related injuries.

LABORATORY ANIMAL MEDICINE

 II. Biology
C. Genetics
Both C. japonica and Gallus domesticus, the common
chicken, have a diploid number of 78 chromosomes,
most of which are orthological (Guttenbach et  al.,
2003); numerous authors have concluded that quail
and chicken lines diverged 35–46 million years ago
(Jiang et  al., 2010; Kan et  al., 2010; Kayang et  al., 2006;
Van Tuinen and Dyke, 2004; Van Tuinen and Hedges,
2001). The International Chicken Genome Sequencing
Consortium has determined that chickens possess
five pairs of macrochromosomes (>40 Mb), five pairs
of intermediately sized chromosomes (20–40 Mb), and
28 pairs of microchromosomes (<20 Mb) (Hillier et  al.,
2004; Wallis et  al., 2004); Japanese quail chromosomes
are similar in this regard (Kawahara-Miki et al., 2013).
Chicken microchromosomes comprise about one-third
of the total genome size (Wallis et al., 2004), yet encode
50–75% of all genes (McQueen et al., 1998; Burt, 2002),
replicate earlier in the S phase than macrochromosomes
(McQueen et al., 1998), and have higher recombination
rates than macrochromosomes (Wong et  al., 2004). It
has been hypothesized that microchromosomes initi-
ated as fragments of ancestral macrochromosomes, and
conversely that macrochromosomes formed by aggre-
gation of microchromosomes 100–250 million years ago
(Fillon, 1998). In contrast to mammals, avian sex chro-
mosomes are designated Z and W; males are ZZ and
females are ZW and thus the ovum governs the sex of
the offspring. There are no mutual genes between the
avian ZW and mammal XY chromosomes, nor do the
ZW and XY chromosomes seem to share a common
origin (Stiglec et al., 2007).
Recently, the female Japanese quail genome was
sequenced using Illumina next-generation sequencing
platforms and assembled against the chicken reference
genome (Kawahara-Miki et  al., 2013). The Japanese
quail genome is ~1.41  Gb in size (Kawahara-Miki
et al., 2013; Nakamura et al., 2004; Tiersch and Wachtel,
1991). There are currently ~150 microsatellite markers
identified for the Japanese quail based on the genome
sequences (Kawahara-Miki et  al., 2013; Tadano et  al.,
2014). The Japanese quail genome sequence and associ-
ated resources will offer essential genetic and genomic
reference information, molecular tools for making
improved primers and nucleic acid probes to use in
polymerase chain reaction, multiplexed hybridization
chain reaction-based RNA in situ, siRNA- and shRNA-
based RNA inactivation, and CRIPR-Cas9 targeting
approaches.
Considerable breeding in laboratory and commer-
cial settings is carried out with quail lines that have
been selected or screened for a particular phenotype
or genotype, e.g., feather coloration, growth rate, mutant
lines, transgenic lines. The most common method used
LABORATORY ANIMAL MEDICINE
1093
to characterize the resulting phenotype is to observe
the hatchlings for feather color, developmental abnor-
mality, etc.
For genotypic analyses, the mesoderm-derived cho-
rioallantoic membrane (CAM), a highly vascularized
membrane in close contact with the pores of the egg-
shell to facilitate gas exchange, can be used to isolate
and analyze the hatchling’s genomic DNA. For example,
transgenic founder lines that express markers such as
fluorescent proteins or lacZ can be initially screened for
transgene expression in the CAM cells or shell membrane
cells of freshly hatched eggs, assuming the transgene is
expressed by the cells being CAM cells. For instance,
a ubiquitously expressed transgene or vascular-specific
transgene (Sato et al., 2010) is readily seen in the CAM
cells when observed using a epifluorescence microscope;
however, a transgene whose expression is restricted to
the glial or adult kidney cells would not be expected to
be visualized in CAM cells. Feather blood cells can also
be visualized using a fluorescent microscope to assay
transgene expression; in addition, genomic DNA can
be analyzed by conventional recombinant DNA tech-
niques to confirm a desired gene or transgene. Southern
blot analysis will determine transgene copy number and
genomic integration site (Sato et al., 2010).
Japanese quail are very susceptible to inbreeding
depression (Sittmann et al., 1966), which reduces popu-
lation fitness via decreased hatchability, viability, and
fertility. Inbreeding increases genome homozygosity
and preserves recessive mutations carried by a popula-
tion (Frankham et al., 2002; Keller and Waller, 2002). The
inbred lines of quail that Sittmann et al. (Sittmann et al.,
1966) initiated by consecutive full-sibling matings only
survived to the third generation. In another study, only
five out of 17 full-sibling mating inbred lines survived
to the fifth generation (Kulenkamp et  al., 1973). Using
this standard approach, there is currently no record of
any inbred Japanese quail line living past eight genera-
tions of consecutive full-sibling matings (Okimoto, R.,
University of Arkansas, personal communication to KC;
Wada, M., Tokyo Medical and Dental University, per-
sonal communication to KC).
As in mice, the increase in genome homozygosity
of inbred quail lines is useful for genome analysis and
gene mapping (Hoti and Sillanpaa, 2006). At the Avian
Research Centre at the University of British Columbia, a
more relaxed inbreeding scheme was employed to gen-
erate an inbred Japanese quail line in order to estimate
the inbreeding coefficient of a quail population and com-
pare it to the inbreeding level that would be attained
with full-sibling mating. The inbreeding scheme entailed
mating full sibling sisters to full sibling brothers from
a different (but related) family (Kim et  al., 2007). This
quail line has now been inbred for 24 generations and
still appears vigorous.
1094 22.  Japanese Quail as a Laboratory Animal Model

III.  LABORATORY MANAGEMENT The type of housing system selected should take into
AND HUSBANDRY consideration the anticipated use of the birds, as well as
regulatory and animal welfare requirements. Housing
A.  Sources of Quail design should accommodate ease of sanitation and
maintenance.
Fertilized quail eggs can be ordered from a com- Outdoor aviaries that provide a semi-natural envi-
mercial supplier. However unlike for chickens, there ronment encourage natural behaviors such as exercise,
is no vendor for particular quail strains with a known flight, social interaction, foraging and dust bathing
breeding history. Quail obtained from commercial farms (Schmid and Wechsler, 1997); access to ample vegetative
may be heterogeneous genetically and may differ from
batch to batch; this situation makes comparisons diffi-
cult between different experiments (Cheng and Nichols,
1992). Researchers desiring a specialized quail model
usually obtain the birds from the laboratory that devel-
oped the model; however, two technical problems can be
associated with such an acquisition. First of all, stringent
and expensive quarantine procedures are required if the
shipment has to cross national boundaries. Occasionally,
there is also a tariff imposed. Importation regulations
are country dependent and typically focus on avian-
transmitted diseases such as avian influenza and sal-
monellosis, among others. It is hoped that adopting the
practices used by commercial avian producers, such
as the flock certification program provided by the US
National Poultry Improvement Plan (NPIP), will facili-
tate the international transfer of eggs and birds.
Second, many specialized quail strains were not
maintained beyond the researcher’s tenure (Fulton and
Delany, 2003), as maintaining bird colonies is expensive
and budget  allocation for conserving a population is
not an administrative priority. Although until recently FIGURE 22.4  Commercial battery-cage housing Japanese quail.
there was no practical procedure for the cryopreserva-
tion of avian germplasm, Liu and colleagues (Liu et al.,
2010, 2012a,b, 2013a–d) have pioneered such a proce-
dure for Japanese quail gonadal tissue that shows great
promise.
It is critical to use freshly laid eggs that come from
a productive and healthy flock. Egg storage and ship-
ping conditions can also contribute to egg fertility, e.g.,
low egg fertility levels may be observed if the eggs are
exposed to high temperatures during transportation;
transportation in cooled containers can mitigate this con-
cern. Eggs should be stored immediately upon receipt in
a refrigerator that is kept at ~13°C and humidified with
an open tray of water; in contrast a 4°C refrigerator is
too cold and will cause embryonic death.

B.  Housing Systems

Housing systems for adult Japanese quail include
commercially available poultry battery cages (Gerken
and Mills, 1993) (Fig. 22.4), custom-built indoor floor
pens (Gerken and Mills, 1993) (Fig. 22.5) and semi- FIGURE 22.5  Custom-built indoor floor pen housing Japanese
natural outdoor aviaries (Schmid and Wechsler, 1997). quail chicks.

LABORATORY ANIMAL MEDICINE

 III.  Laboratory Management and Husbandry
cover is critical. In general, key features of a semi-natural
environment are shelter from weather extremes, protec-
tion from predators, and ease of sanitation, egg collec-
tion, and capture. Similar to outdoor aviaries, indoor
floor pens can promote the expression of natural behav-
iors and animal welfare through the provision of artifi-
cial cover and opportunities for exercise, foraging, dust
bathing, and social behavior (RSPCA, 2001). Resource
competition for food and water, as well as cover and
dust baths, must be considered when housing quail
in floor pens; provision of resources must be adjusted
based on stocking density. Battery cages with sloped
floors are widely used in commercial applications; these
facilitate egg collection but result in an impoverished
environment, i.e., decreased opportunities for exercise
and other species-specific behaviors. The percentage of
foot lesions, bone fractures, and beak deformities, as
well as head and eye injuries resulting from aggressive
pecking behavior, were increased in birds housed in bat-
tery cages compared to birds housed in deep litter floor
pens (Abdelfattah et al., 2012). Because the same authors
found that birds housed in deep litter floor pens demon-
strated significantly increased cortisol levels and mortal-
ity attributed to increased disease, including parasitism,
consideration should be given to increased health moni-
toring for infectious agents in quail housed in situations
where routine sanitation may be difficult and substrates
may remain damp for periods of time. The NRC Guide
for the Care and Use of Laboratory Animals (2011) lists
minimum space recommendations for quail as 0.25 feet2
floor area/animal. Increased cage space resulted propor-
tionally in significant improvement in body weight gain,
age at 50% egg production, egg production, food conver-
sion values, and decreased mortality (Nagarajan et  al.,
1991). Overall, aviary room temperatures of 22–25°C are
considered ideal for quail.
It has been reported that the typical vertical escape
response in quail may lead to serious head injuries if
cage ceilings are too high; one study concluded that cage
heights should be less than 25 cm to reduce head injuries
associated with the escape response, but at least 20 cm
to allow birds from heavy strains to stand up fully in a
normal postural position (Gerken and Mills, 1993). In
contrast, other experts (RSPCA, 2001) suggest that cages
of 30 cm in height reduced or eliminated the problem of
head injuries in quail; another proposed solution is cage
roofs comprised of materials that do not cause head inju-
ries, e.g., a flexible plastic grid. Because upward flight is
part of the normal behavior repertoire for quail, further
study into the topic of cage height would be useful.
Environmental enrichment and habituation to human
presence and environmental change can serve as ame-
lioration strategies for panic reactions in quail (Mench
and Blatchford, 2014).
LABORATORY ANIMAL MEDICINE
1095
Japanese quail may be housed on a number of dif-
ferent types of bedding substrates including wood
shavings, chopped corn cobs, ground bark, and peanut
hulls. Use of small, solid-bottom cages coupled with
the type of food (i.e., wheat-based high gluten content)
and bedding provided may result in accumulation of
food on the digits (referred to as toe-balls) which may
lead to trauma to the digits if left untreated. It should
be appreciated that bedding substrates can be a source
of contamination.
C.  Food and Water
Food and water are typically provided ad libitum to
quail; the delivery method for both is dependent on the
type of housing and the age of the birds. Food receptacle
design should encourage easy access while preventing
the birds from soiling uneaten portions; for example,
birds in cages with food dishes will often nest in the
feeders, resulting in spillage and soiling. Placement of
feed troughs outside of the cage reduces spillage, pre-
vents soiling of the food, and facilitates ease of food
provision. Trough feeders and waterers placed outside
of the cage are commonly used in battery cages. Birds
tend to actively investigate water sources, and cage
floods can occur if the water source is placed inside a
solid floor cage or pen. Quail readily adapt to use of an
automatic drinking valve with an integral cup basin,
whereas gravity-fed bell drinkers are commonly used
for newly hatched chicks and juveniles. Grill screens,
rubber rings, or pebbles should be used to reduce the
available water drinking area for very young chicks, as
otherwise they may drown (Randall and Bolla, 2008).
Drinking devices should be inspected daily to ensure
that they are working properly and not clogged with
debris. Water consumption varies with age and strain,
with 2-week-old chicks ingesting 23.3 ml daily, 4-week
old birds drinking 30.0 ml per day (Farrell et  al. 1982),
and 7-week-old birds ingesting 40.9–62.1 ml/g per day
(Visser et al., 2000; Cheng et al., 2010).
Wild quail are omnivorous, eating primarily seeds and
plant matter but adding terrestrial invertebrates in the
summer (del Hoyo et al., 1994). In captivity, the dietary
requirements for Japanese quail vary depending on the
age and genetic selection for meat or egg production
(Table 22.4). In general, a commercial game bird or tur-
key starter diet with 24.0–26.0% crude protein should be
provided for the first 6–8 weeks after hatching followed
by a game bird or turkey finisher diet that contains less
protein. The pelleted starter diet must be ground into a
powder form for feeding chicks for the first two weeks
post hatch. Facility staff should be aware that diets high
in wheat content may mix with fecal material and stick
to the digits of the birds. As expected, food consumption
1096 22.  Japanese Quail as a Laboratory Animal Model

TABLE 22.4 Diet Specifications for Japanese Quail and efficiency varies by age and gender (Table 22.5).
(as Percentage or Unit per Pound of Diet) Specially prepared diets may be purchased from feed
Starter/
companies, and a synthetic diet has been developed for
grower Finisher Japanese quail used in research (Cheng et al., 2010). For
experimental purposes, high-cholesterol diets have been
>6 weeks to Adult
used to induce atherosclerosis in Japanese quail (Cheng
Nutrient Unit 0–6 weeks market breeder
et al., 1997; Godin et al., 2001, 2003; Hoekstra et al., 2003,
Crude protein % 24.0–26.0 17.0–19.0 18.0–20.0 2004). Game bird or turkey feeds incorporating coccid-
Metabolizable kCal 1315 1315 1315 iostats (i.e., monensin sodium (Coban) and amprolium)
energy and/or antibiotics are commercially available.
Calcium % 1.80 0.70 2.50
Nonphytate % 0.30 0.25 0.35 D.  Common Procedures
phosphorus
Adults and chicks can be individually identified
Sodium % 0.15 0.15 0.15
through the application of commercially available leg
Methionine % 0.50 0.42 0.45 bands or wing tags. Plastic coil legs bands (Size 4)
Methionine + % 0.75 0.68 0.70 obtained in multiple colors (National Band and Tag
cystine Company, Newport, Kentucky, USA; www.national-
Lysine % 1.30 0.90 1.00 band.com) can be used to identify chicks the day after
hatching; however, the maximum diameter of these
Threonine % 1.02 0.85 0.74
bands precludes their continued use once the birds
Tryptophan % 0.22 0.20 0.19 reach 2–3 weeks of age. Colored aluminum leg bands
Percentage amount per lb of diet in size 8 can be applied after the birds are 3 weeks of
age. Wing tags are suitable for birds of all ages, but
Vitamins added per lb
of diet 100% 80% 100%
proper application in the propatagium of newly hatched
chicks is difficult due to their small size. The wing tags
Vitamin A IU 3000 must be positioned just behind the tendon along the
Vitamin D ICU1 1000 leading edge of the wing, opposite the humeral–ulnar
Vitamin E IU 18.0 joint. Improper placement can result in traumatic dam-
age to the musculature of the wing as the bird develops.
Vitamin K mg 1.0
Other recommendations are for the use of #5 fingerling
Thiamin mg 1.0 tags (National Band and Tag Company) in hatchlings,
Riboflavin mg 2.8 and aluminum chick wing tags for adult birds. Rodent
ear tags can also work as wing bands in chicks and
Niacin mg 20.0
adult birds.
Choline mg 115 Capture of caged Japanese quail can be achieved with
Pyridoxine mg 1.5 relative ease; it should be performed calmly but quickly
Pantothenic acid mg 7.0 to avoid injuries during escape attempts. Chicks should
be held carefully in the palm of the hand, using the
Folic acid mg 1.0
thumb and forefinger for restraint. Adult birds should
Vitamin B12 μg 5.0 be restrained by pinning the wings against the body
Biotin μg 50.0 while allowing the legs to hang loose. Alternatively, the
bird may be gently cupped around the wings using both
Trace minerals added per lb of diet
hands. Attempts to restrain the bird by the legs can result
Manganese mg 25.0 in traumatic injury to the legs, and failure to restrain the
Iron mg 30.0 wings may result in damage to the wings, including
Copper mg 5.0
bone fractures. Capture of birds housed in large pens or
aviaries usually requires netting of the birds.
Zinc mg 30.0
When the suitability of five different sites (brachial,
Iodine mg 0.2 jugular, caudal tibial vein, external dorsal thoracic vein
Selenium mg 0.136 and the heart) was compared for blood sampling and
intravenous injections in Japanese quail, the jugular vein
From http://www.aces.edu/pubs/docs/A/ANR-1343/index2.tmpl Alabama Cooperative
Extension System ANR-1343. was reported to be most successful (Arora, 1979). Small
1
ICU = International Chick Unit. volumes of blood from the ulnar vein may be collected
in microcapillary tubes.

LABORATORY ANIMAL MEDICINE

 IV. Breeding and Rearing Quail 1097
TABLE 22.5 Body Weight, Cumulative Feed Consumption, and Feed
Efficiency of Japanese Quail

Body weight (g)1 Feed consumption (g) Feed efficiency (g:g)2

Age
(weeks) Male Female Male Female Male Female

2  40  40  50  50 1.25 1.25

4  90 100 180 190 2.00 1.90
6 120 130 300 330 2.50 2.53
8 130 160 350 450 2.69 2.81
10 140 170 400 510 2.86 3.00
From http://www.aces.edu/pubs/docs/A/ANR-1343/index2.tmpl Alabama Cooperative Extension System
ANR-1343.
1
To convert gram values to pound units, divide by 454.
2
Feed efficiency is the amount of feed required per unit of body weight.

Similar to other small avian species, Japanese quail 100

can be anesthetized by exposure to an inhaled gas anes-
thetic (e.g., isoflurane) or through the administration of
injectable anesthetics. Oral or intramuscular adminis-
tration of ketoprofen at 2 mg/kg resulted in very low
bioavailability and the drug exhibited a very short half-
life in Japanese quail (Graham et  al., 2005). Changes in
complete blood count and serum chemistry were mini-
mal but injection site muscle necrosis was observed in
Japanese quail following intramuscular administration
of 2 mg/kg meloxicam every 12 h for 14 days (Sinclair
et al., 2010). Japanese quail can be euthanized by expo-
sure to inhaled anesthetic gas (e.g., isoflurane) or CO2;
chicks and embryos that are beyond 50% incubation are
acclimated to high CO2 concentrations in the egg and
thus require prolonged exposure to CO2 for euthanasia.
A high dose of an injectable anesthetic is also an effec-
tive method of euthanasia. Physical methods, including
cervical dislocation (between the first and second verte-
brae) or decapitation, are acceptable on the condition that
personnel are properly trained and proficient in humane
application of these techniques. Prolonged exposure to
CO2, cooling (<4°C for 4 h), or freezing is recommended
for eggs at less than 50% incubation (AVMA, 2013).
IV.  BREEDING AND REARING QUAIL
A. Reproduction
Reproduction in Japanese quail is strongly influ-
enced by the photoperiod (Robinson and Follett, 1982;
Yasuo et al., 2006), such that wild quail breed in spring
and summer in response to increasing day length.
When housed under artificial light with a day length
of 12 h or more, Japanese quail will breed year round.
Photoperiod influences sexual behavior, cloacal protru-
sion and cloacal foam production (Sachs, 1969). Changes
80
Egg laying rate (%)
60
50
40
20
0
0 5 10 15 20
Age (month)
FIGURE 22.6  Egg production as a function of age in domestic
Japanese quail. Data from Minvielle et al. (2000).
in photoperiod result in neuroendocrine changes in both
male and female quail (Ball and Balthazart, 2010), and
short day lengths result in gonadal regression and breed-
ing cessation (Follet and Pearce-Kelly, 1990).
In general, Japanese quail reach sexual maturity at
6–8 weeks of age, but this is dependent on photoperiod
and strain. Young quail raised under short-day lighting
schemes, which mimic fall and winter days, either do not
mature sexually or show delayed maturation (Delville
et al., 1984). Lighting regimes for commercial producers
vary but are typically designed with 16 h of light per
day (Gerken and Mills, 1993). Hens typically begin to
lay eggs consistently at ~8–10 weeks of age, with egg
fertility rates quickly increasing over time; with time,
egg production gradually declines (Fig. 22.6). Japanese
quail exposed to continuous light gained weight faster
and reached sexual maturity earlier than birds housed

LABORATORY ANIMAL MEDICINE

1098 22.  Japanese Quail as a Laboratory Animal Model
under 13 h of daylight (De Jager, 2003). Female puberty,
characterized by the average age of the first egg laid in
a flock of Japanese quail maintained on a 16:8 day: night
cycle, occurred at 48.8 days of age, while sexual matu-
rity, characterized by the time at which egg production
for the flock reached 50%, occurred at 54.2 days. Puberty
in the male, defined by the first appearance of secondary
sex characteristics like crowing and production of cloa-
cal foam, occurred at 32.6 days of age; in contrast, sexual
maturity in the cock, or the appearance of mating behav-
ior and pronounced foam gland secretion, occurred by
42.4 days of age (Abdelfattah et  al., 2012). Maximum
fertility in age-matched male and female group-housed
birds maintained under a 16:8 day: night cycle was
reached during the fourth month of age and then gradu-
ally declined (Abdelfattah et al., 2012). Fertility in battery
cage-housed Japanese quail is also influenced by the
ratio of males to females. While a male: female ratio of
1:3 provided optimal fertility, a male: female ratio of 1:5
resulted in a significant decrease in fertility (Abdelfattah
et  al., 2012). Fertility was also found to be significantly
increased in birds housed in deep litter floor pens com-
pared to birds in battery cages.
B.  Egg Storage and Incubation
The average Japanese quail egg weighs 10–11 g, which
corresponds to about 8% of the hen’s body weight. Egg
size, shape, and color patterns vary considerably based
on the parent’s strain; they are typically tan with brown
speckles, but can also appear snowy white, mottled
brown with a chalky blue covering, or other (Fig. 22.7).
Each laying hen produces 10–12 eggs every 2 weeks
year round in captivity under controlled light cycles.
Wild populations typically lay eggs from late April to
FIGURE 22.7  Japanese quail eggs exhibiting variation in size,
shape, and color patterns.
LABORATORY ANIMAL MEDICINE
early August in Russia, and from late May to August in
Japan. Clutch sizes also vary, with 9–10 eggs per clutch
in Russia and 5–8 in Japan. The female incubates the
eggs in the wild (del Hoyo et al., 1994); however, in cap-
tivity when males are co-housed with females, persistent
male copulatory behavior prevents the hens from brood-
ing (Cain and Cawley, 1972).
In the laboratory setting, quail eggs should be col-
lected and stored daily. Egg collection is best performed
early as most eggs are laid in the late afternoon or eve-
ning. Identification numbers can be recorded on the
eggshell in permanent ink to facilitate tracking of egg
numbers and fertility rates for different transgenic lines.
As eggs are often soiled during egression, they should
be washed with lukewarm water to remove debris and
then air dried. Bacterial penetration across the eggshell
is dependent on bacterial survival on the eggshell sur-
face and egg storage conditions (Gole et al., 2014). Once
dry, eggs should be transferred to commercially avail-
able egg cartons designed to accommodate quail eggs.
Fertilized quail eggs are best stored with their large end
up or on their sides; these recommendations improve
embryo development and orient the embryo in an exper-
imentally desired position. Ideally, eggs should be stored
in a refrigerator set to 13°C and equipped to provide 65%
relative humidity. Turning eggs in their egg flats once
daily improves hatchability. Eggs may be stored at ~11°C
for up to 15 days, 21°C for up to 10 days, and 27°C for up
to 5 days without a significant decrease in hatchability
(Garip and Dere, 2011). Prior to incubation, refrigerated
eggs should be maintained at room temperature until
condensation on the shell has dried.
To optimize egg development, a quality egg incuba-
tor is essential. Many types and sizes of incubators can
be purchased, and the best choice for a given situation
will depend on the extent of planned experiments along
with cost considerations. A forced-air incubator that uti-
lizes an internal fan to circulate air inside the chamber
is ideal. The egg incubator should include an automatic
egg-turning option that is capable of tilting the egg trays
through a 90° angle once an hour (Fig. 22.8). A sepa-
rate hatching incubator equipped with hatching trays
and matching wire mesh lids will be needed to hatch
the eggs. Digital thermostats allow steady, consistent
temperatures and humidity levels to be maintained in
modern incubators. The incubator’s internal tempera-
ture and relative humidity should be monitored at initial
setup and daily during incubation with a high-quality,
calibrated thermometer. Keeping a stock of numerous
replacement parts is recommended to facilitate timely,
in-house repairs. Incubators should be connected to
electrical circuits with generator backup to ensure that
power spikes or outages do not damage the eggs.
An incubation temperature of 38°C and 50–65% rela-
tive humidity results in optimal embryonic development.
 IV. Breeding and Rearing Quail
FIGURE 22.8  Commercial poultry incubator. Reproduced with per-
mission of GQF Manufacturing Company.
The egg trays should be slowly rotated up to a total of
90° every hour until ~E13 to facilitate normal develop-
ment. Eggs may be candled to inspect embryo viability
using a standard commercial egg candler; however, cau-
tion is required to avoid overheating. Although staging
early quail development by candling is difficult due to
the blotchy colored eggshells (Fig. 22.6), viability can
be reliably determined after about 7 days of incubation.
Typical fertility ranges are from 90 to 95%, and peaks in
embryonic mortality may occur during the first 3 and
last 2 days of the 16–18 days of incubation.
Of interest is that quail embryos and chicks commu-
nicate with one another prior to hatching. This behavior
is coincident with the onset of breathing, when quail
embryos begin to acoustically communicate using click-
ing sounds which helps them synchronize their hatching
times (Vince and Salter, 1967; Vince, 1964a,b). The same
authors also reported that clicking sounds can acceler-
ate or decelerate embryonic development and hatching
times (Vince, 1966, 1968a,b, 1973; Vince et al., 1984).
C. Hatching
On the 16th day of incubation, quail eggs should be
transferred to an egg hatching incubator (Fig. 22.9), or
hatchers, and placed on their sides in trays or compart-
ments; the hatchers must be maintained at ~38°C and
~70% relative humidity. Trays of water should be placed
in the incubator to maintain a high humidity, which
will keep the chicks from adhering to their shell mem-
branes during the hatching process. Relative humid-
ity, the ratio of the current absolute humidity to the
LABORATORY ANIMAL MEDICINE
1099
FIGURE 22.9  Commercial poultry hatching unit. Reproduced with
permission of GQF Manufacturing Company.
highest possible absolute humidity (which depends on
the current air temperature), can be measured using a
wet bulb hygrometer. The trays should be covered with
soft, absorbent paper to provide good traction for the
hatchlings; if the lining material is too smooth or slick,
hatchlings often develop splay leg. If the wire mesh is
too large, the hatchlings can fall through or get their
heads, wings, or legs stuck in the wire mesh. Once the
hatchlings start to pip out the shell, the hatcher should
be kept closed so the humidity levels remain high
(~60–70%). Hatchling quail will typically emerge from
their shells over the course of several hours.
In some cases, it may be important to keep hatchlings
separated from one another if they are derived from
distinct genetic or transgenic lines. If so, inexpensive
separators can be made of plastic, cardboard, or other
disposable material, and placed in the hatching trays to
create separate cubicles. The design of hatching cubicles
(each ~10 × 10 cm) can be such that each compartment
has space for a single egg and its chick. This keeps the
hatchling in the same container as its egg to ease phe-
notypic and genotypic analysis of both. Quail eggs lose
approximately 13% of their 10- to 11-g starting weight
via evaporation prior to hatching. The hatchlings typi-
cally weigh about 6–8 g and are brownish with yellow
stripes. During the first 24 h in the hatcher, no food or
water is necessary because the hatchlings obtain nutri-
tion from their yolk sac that retracted into their coelom
in the days prior to hatching. Understandably, hatched
chicks are exhausted, but they typically gain strength
and increase activity over 24 h. Within 24 h of hatching,
the chicks should be transferred to a separate brooder
1100 22.  Japanese Quail as a Laboratory Animal Model
that provides an auxiliary heat source during the first 4
weeks of life.
D.  Rearing Chicks
Poultry brooders suitable for Japanese quail can be
obtained from multiple commercial sources. The brooder
floor should be lined with absorbent paper for the first 2
weeks to prevent leg injuries to the newly hatched chicks.
Game bird or turkey starter diet should be ground into
a powder form and sprinkled around the brooder floor
for the hatchings to eat. Unrestricted access to water
may be supplied through use of small bell jars fitted
with water troughs; size appropriate glass marbles must
be placed in the water troughs to prevent the hatch-
lings from falling into the water well and drowning. The
temperature inside the brooder should be maintained
at 37°C for the first week post-hatch. The heat source
should be placed such that temperature gradients are
established and chicks can select their desired tempera-
ture zone. Chick behavior should be observed daily or
more frequently, and the temperature adjusted if they
are huddled under the heat source (suggesting the heat
source may be inadequate) or along the outer perimeter
of the brooder (suggesting the heat source is providing
too much warmth). The brooder temperature should be
decreased 2–3°C every week for three weeks (Hodgetts,
1999). At 2 weeks post-hatch, the paper liner should be
changed to a bedding substrate (paper chip, wood chip,
or other suitable material). At 4 weeks post-hatch, the
birds no longer need an external heat source. The diet
should be changed to a game bird or poultry finisher/
breeder diet when the birds are 5 weeks post-hatch.
V.  DISEASES/WELFARE CONCERNS
A. Noninfectious
Japanese quail are susceptible to many common
poultry diseases. Detailed reviews of diseases affecting
quail, along with information on control, prevention,
and treatment have been published (Barnes, 1987; Reed
and Jack, 2013). As for other species, the implementa-
tion of biosecurity measures is critical to prevent intro-
duction of infectious agents into a quail colony. Such
measures include sanitation and quarantine practices;
effective insect and rodent control programs; and the
inspection of food, water, and bedding materials for
potential contamination. A risk of disease transmission
is reduced by the use of fumigated eggs rather than
adult birds. The health status of the source flock should
be carefully evaluated prior to transfer of eggs or birds.
Ideally, egg incubation and hatching areas are physically
separated from other areas, and human traffic always
LABORATORY ANIMAL MEDICINE
flows from clean to dirty areas. Eggs from an external
source should be carefully washed using a quaternary
ammonium (250 ppm) solution and water temperature
of 43–49°C. Washed eggs should be kept apart from
soiled eggs, and egg flats should be cleaned and disin-
fected prior to use. A comprehensive sanitation program
should be implemented for incubators and brooders as
well as for the housing area for adult birds. Bedding
substrate, food, and water must also be considered as a
possible contamination source and should be evaluated
and/or treated to reduce risk of pathogen transmission.
Personnel with access to the housing facilities should not
have contact with other birds for 72 h. A health surveil-
lance program should also be implemented.
Trauma from conspecific pecking is one of the most
frequently observed health concerns in laboratory colo-
nies of Japanese quail. Head injuries from pecking are
typically located toward the back of the head and neck
areas (Fig. 22.10), while head injuries from bumping
the cage ceiling are located more toward the top of the
head. As these injuries commonly result from aggressive
pecking in adult male quail, multi-male breeding groups
are not generally recommended (Wechsler and Schmid,
1998). However multi-male breeding groups may be
effective if (1) the male: female sex ratio is not too low
(e.g., not less than 1 male:3 females), (2) the enclosure
is adequately sized, (i.e., no high-density rearing), and
(3) there are structures in the enclosure where birds can
hide. A ratio of 1 male to 3 females is considered optimal
in small, confined spaces. Even so, breeding groups with
more space should be limited to a single male plus two
to five females, as fertility will decrease if the sex ratio
is higher than 1:5. Regardless, pecking injuries may still
occur in single male breeding groups when females and
males peck one another. In intensive breeding situations
FIGURE 22.10  Head injury caused by aggressive pecking by cage
mates.
 V. Diseases/Welfare Concerns
with limited space and hiding areas, females may exhibit
feather loss and occasional wounds to the back and head
related to male mounting behavior. Misdirected copula-
tion attempts by the male may result in head wounds
and eye trauma of the female.
Depending on the anatomical location and severity
of the trauma, the injured bird should be transferred to
a separate enclosure and administered an anti-inflam-
matory drug such as ketoprofen (2–5 mg/kg IM). Birds
with mild injuries can often be returned to the mating
group once the injuries have healed. For birds raised in
a laboratory setting, trimming the beak and nails with a
conventional nail clipper every 4 weeks can reduce the
number of pecking related-injuries. Kwik-Stop styptic
powder (ARC Laboratories, Atlanta, GA) is effective in
stopping bleeding that occurs coincident with beak and
nail trimming. Alternatively, beak trimming can be per-
formed through the use of a poultry hot-blade beak trim-
mer, which has an adjustable guard to keep from cutting
too deep and also seals off blood vessels. However, hot-
blade beak trimming is controversial due to the possibil-
ity for acute and chronic pain, as well as possible beak
re-growth (Cheng, 2011). Infrared beak trimming per-
formed on 1-day-old domestic chickens has been shown
to be more effective at reducing beak re-growth and is
associated with fewer side effects compared to hot-blade
beak trimming (Dennis and Cheng, 2010). Animal wel-
fare is enhanced when birds are housed in floor pens on
natural substrate, as normal behaviors can negate the
need for beak and nail trimming.
Foot injuries can be observed in battery-housed or
deep-litter-housed quail. When housed in solid bottom
cages or pens and fed a wheat-based (high gluten) diet,
secretion from the male foam gland mixes with fecal
matter and food and may adhere to the toes of birds
FIGURE 22.11  Accumulation of fecal material, food, and male
cloacal gland foam on digits.
LABORATORY ANIMAL MEDICINE
1101
(Fig. 22.11). Once dried, the adhered material is often dif-
ficult to remove without injuring the bird’s toes and nails.
Pododermatitis in battery-caged quail (Abdelfattah et al.
2012) may also be associated with trauma due to abra-
sions caused by the wire bar floor. Provision of a solid
resting surface can reduce foot lesions in battery-housed
quail; nevertheless, scheduled monitoring of quail feet
should be part of a routine health monitoring program
established for Japanese quail.
Few neoplasms have been reported in Japanese quail.
Sertoli cell tumors were reported in 9% of 3- to 5-year-
old males kept in a laboratory colony for aging stud-
ies (Gorham and Ottinger, 1986). Leiomyomas were
reported in the oviduct ligaments of rapidly growing
Japanese quail (Foster et al., 1989; Tuzcu et al., 2010).
B. Infections
1. Viruses
Similar to other gallinaceous birds, Japanese quail are
susceptible to a number of avian viral pathogens, most
commonly quail bronchitis virus (QBV), Newcastle virus,
and Eastern Equine Encephalomyelitis (Shivaprasad,
2002). Although a brief description of viral pathogens
in Japanese quail is provided below, the reader can refer
to more comprehensive information elsewhere, e.g.,
Swayne et al. (2013).
QBV, a type I avian adenovirus, has been documented
to cause clinical disease in Japanese quail (Reed and
Jack, 2013). The route of transmission is uncertain but
likely via aerosolization. QBV is highly contagious in
susceptible flocks, resulting in rapid morbidity and mor-
tality; severe illness is most often observed in chicks of
less than 6 weeks of age. Clinical signs include decreased
appetite, ruffled feathers, open-mouth breathing, rales,
sneezing, nasal and ocular discharge, and death. Older
birds may remain asymptomatic. Gross necropsy find-
ings include tracheal opacity, thickening of the trachea
causing partial obstruction, and the presence of muco-
sal exudate. Histopathologic lesions are characterized
by large basophilic intranuclear inclusions; necrotizing
proliferative bronchitis; and hepatic, splenic, and cloacal
bursa necrosis.
Newcastle disease, caused by avian paramyxovirus
I, can infect Japanese quail (Usman et al., 2008). Clinical
signs range from subclinical infection to lethargy, ruffled
feathers, dyspnea, torticollis, paralysis, and hemorrhagic
diarrhea. Gross lesions include tracheal hemorrhage,
inflamed and hemorrhagic Peyer’s patches, pulmonary
edema, hemorrhage of the proventriculus, and intestinal
congestion. Transmission is horizontal via the fecal–oral
route or by inhalation of contaminated dust.
Japanese quail are susceptible to Eastern Equine
Encephalomyelitis, which is caused by an arbovirus
1102 22.  Japanese Quail as a Laboratory Animal Model
(Alphavirus, family Togaviridae); the virus may be
transmitted by arthropod vectors, primarily the mos-
quito, Culisetta melanura, or through feather picking and
cannibalism. Infected birds exhibit depression, tremor,
paralysis, torticollis and death. A primary gross find-
ing in a commercial quail flock was duodenal catarrhal
enteritis (Eleazer et al., 1978).
Duck Adenovirus A, also known as egg drop syn-
drome, has been reported in Japanese quail; it can be
intermittently shed in feces as well as vertically trans-
mitted, with viral particles on the external surface of
the egg and internally. Clinical signs of decreased egg
production, decreased eggshell pigment, and the pro-
duction of thin or soft-shelled eggs have been described.
To date, this virus has not been reported in the United
States.
Hydropericardium syndrome in a commercial flock of
Japanese quail has been attributed to infection with fowl
adenovirus serotype 4 (FAV-4). Infection resulted in 4%
mortality and was characterized by hydropericardium
and mild-to-moderate hepatomegaly accompanied by
splenic and kidney congestion (Roy et al., 2004).
Natural outbreaks of Marek’s disease, a lymphop-
roliferative disease caused by a cell-associated herpes
virus, are relatively common in commercial flocks of
Japanese quail (Nagarajan et  al., 2013). Transmission
occurs through direct contact, and mortality in unvac-
cinated flocks may range from 10–20%. Gross lesions are
observed in the proventriculus, spleen, and liver. Cooper
et al. (2007) have presented a case report of Marek’s dis-
ease in laboratory-housed Japanese quail. Clinical signs
of disease included lethargy, anorexia, weight loss, soft
feces, and lime-green urates.
Japanese quail have been shown to be more suscep-
tible to infection with avian influenza virus than turkeys
(Bonfante et  al., 2013); quail were readily infected with
lower challenge doses and transmitted the virus to other
birds without showing signs of clinical disease. Major
influenza outbreaks in quail are uncommon (Perez et al.,
2003).
Natural infection with quail poxvirus has been
reported in a commercial egg-laying flock of Japanese
quail (Gulbahar et  al., 2005). Clinical signs of disease
included weight loss, blepharitis, conjunctivitis, blind-
ness, crusty papules at the commissures of the beak and
around the external nares, decreased egg production,
and impaired fertility. Infection was associated with 60%
morbidity and 20% mortality. Quail pox is a distinct spe-
cies of the genus Avipoxviridae, and vaccination with
pigeon or fowl poxviruses does not provide protective
immunity to quail.
Japanese quail are susceptible to avian encephalo-
myelitis virus, a member of the Picornaviridae family.
Naturally occurring infection results from fecal–oral and
LABORATORY ANIMAL MEDICINE
vertical transmission. Clinical signs present in young
birds, typically 1–2 weeks of age, include ataxia and trem-
ors, especially of the head and neck. Morbidity among
chicks is typically 40–60%. Depressed egg production,
decreased hatchability and increased embryo mortality
can be present in hens exposed as adults. Virus may be
shed from 5 days in adult birds and up to 2 weeks in
young birds. Gross necropsy lesions are often limited to
whitish areas in the muscularis layer of the ventriculus,
as a result of infiltrating lymphocytes. Histologic lesions
include a disseminated, nonpurulent encephalomyelitis,
ganglionitis of the dorsal root ganglia, and hyperplasia
of lymphoid follicles in the proventriculus, ventriculus,
and myocardium.
Reticulendotheliosis virus (REV) is a retrovirus found
in a variety of domestic and wild birds; Japanese quail
comprise a natural host for REV. Viral transmission
occurs through direct contact. REV infection can result
in immunosuppression, runting syndrome, high mortal-
ity, acute reticulum cell neoplasia, or T-cell and/or B-cell
lymphomas. Histopathological lesions in REV infection
are similar to those found in avian lymphoid leukosis
and Marek’s disease. Incidence of disease in commercial
poultry flocks is sporadic.
2. Bacteria
Ulcerative enteritis or ‘quail disease’ is a fatal
enteric disease caused by Clostridium colinum, primar-
ily in captive quail but also in several other avian spe-
cies; genetic susceptibility in Japanese quail may vary
(Collins et al., 1975). Young quail are most susceptible at
4–12 weeks of age, but disease may also occur in older
birds. Transmission is via the fecal–oral route with con-
taminated food, water or litter being the most common
source. Morbidity and mortality often depend on fac-
tors such as concurrent coccidiosis, overcrowding, food
and water withdrawal, medication, and management.
Clinical signs can be acute death with no premonitory
signs, diarrhea, and emaciation. In young quail, 100%
mortality can be seen. Gross necropsy findings include
duodenal hemorrhagic enteritis with mucosal ulceration;
the liver and the spleen may also exhibit pathological
changes. Survivors may be carriers.
Japanese quail can be infected by two serovars of
Salmonella enterica; these are often referred to as pul-
lorum disease (Salmonella pullorum) and fowl typhoid
(Salmonella gallinarum). Distribution is worldwide and,
importantly, there is a potential for zoonotic transmis-
sion. Transmission is both horizontal via the fecal–oral
route and vertical via transovarian infection. Chicks
can present with weakness, decreased appetite, white
chalky material on the vent, respiratory signs, joint
swelling, anemia, and death. Adult birds may be sub-
clinical or can exhibit generalized signs of disease,
 V. Diseases/Welfare Concerns
e.g., decreased appetite, diarrhea, depression, ruffled
feathers, and decreased egg production. Gross lesions
in quail include splenomegaly and hepatic lesions.
Recovered birds are resistant to re-infection but may
remain carriers. The bacterium is susceptible to com-
mon disinfectants.
Campylobacteriosis leads to acute gastroenteritis in
birds, including Japanese quail, and humans (Maruyama
and Katsube, 1988, 1990). It is caused by the gram-neg-
ative, microaerophilic bacteria Campylobacter jejuni and
C. coli. Horizontal and vertical transmission are pos-
sible. Clinical signs may not be apparent or diarrhea
can be present. Gross lesions are typically minimal, and
confined to the gastrointestinal system. In one study,
C. jejuni was reported to propagate in the spleen, liver,
and lungs and to persist up to 17 days in orally inocu-
lated Japanese quail (Maruyama and Katsube, 1988).
Worldwide, colibacillosis associated with avian
pathogenic Escherichia coli infection is the most common
infectious bacterial disease in poultry; the bacteria is
responsible for significant economic losses in the poul-
try industry (Nolan et  al., 2013). Clinical signs include
lameness, anorexia, dehydration, omphalitis, lethargy
and mortality; histopathologic lesions can be localized,
e.g., osteomyelitis, cellulitis, or systemic.
Pasteurella multocida, the causative agent of fowl chol-
era, is a contagious disease affecting a number of avian
species, including quail. Clinical signs of acute disease
may include fever, anorexia, oral discharge, diarrhea,
dyspnea, and cyanosis. Birds with chronic disease may
exhibit swollen joints including footpads, sternal bursae,
wattles or sinuses, torticollis, and signs of respiratory
impairment. Mortality due to acute septicemia has been
reported in a quail colony bred for experimental use
(Goto et al., 2001).
Infection with Bordetella avium infection has been asso-
ciated with a highly contagious upper respiratory tract
disease in Japanese quail; the natural host for B. avium is
the turkey. Clinical signs include sneezing, cough, nasal
exudate, and dyspnea, and gross lesions are typically
confined to the upper respiratory tract.
Naturally occurring infection with Mycoplasma galli-
septicum and M. synoviae has been reported in Japanese
quail. Transmission may be horizontal or vertical. Clinical
signs of M. gallisepticum infection include nasal discharge,
increased lacrimation and sinusitis. Sinusitis and arthritis
has been attributed to M. synoviae infection in quail.
Chlamydiosis is caused by the bacterium Chlamydia
psittaci; clinical signs and lesions in Japanese quail are
similar to other avian species. Transmission results
from inhalation or ingestion of contaminated mate-
rial. Infected birds excrete the agent in feces and nasal
discharge. Chlamydia psittaci is a zoonotic agent and is
transmitted via infectious aerosols.
LABORATORY ANIMAL MEDICINE
1103
Japanese quail are likely susceptible to a number of
other bacterial pathogens, such as Mycobacterium avium,
Staphylococcus spp., Streptococcus, and Listeria. Reports
of naturally occurring disease due to these organisms
in quail are rare.
3.  Protozoa Agents
Coccidiosis, caused by coccidian protozoa, Eimeria
spp., has been diagnosed in Japanese quail (Teixeira
et  al., 2004). In young quail chicks, infection is charac-
terized by weight loss and diarrhea; it can be fatal or
leave the bird with compromised digestion leading to
malnutrition. Sulfonamides are an effective treatment.
Cryptosporidiosis, a diarrheal disease caused by
infection with Cryptosporidium sp. has been reported
in commercial quail colonies. Infected quail may have
respiratory and intestinal involvement similar to chick-
ens. The parasite is protected by an outer shell that
allows survival outside the body for long periods and
resistance to chlorine and many other disinfectants;
however, it is killed by exposure to temperatures greater
than 65°C.
4. Fungi
Infection with fungal agents is rare in quail. In some cases,
transmission has been associated with contaminated lit-
ter and incubators. Aspergillosis caused by infection with
Aspergillis fumigatus results predominantly in pulmonary
disease, and may cause significant morbidity and mortal-
ity in young birds. Crop mycosis from Candida albicans is
the most common fungal infection of the digestive tract.
Zygomycosis and ochroconis are uncommon fungal
infections in poultry. Ochroconis gallopava was reported as
the causative agent in an outbreak of fungal encephalitis
in Japanese quail chicks. Clinical signs were incoordina-
tion, loss of equilibrium, tremors, torticollis, paralysis,
and death; histologically, fungal hyphae were apparent
(Dykstra et al., 2013). Histoplasmosis and cryptoccocco-
sis result in rare infections in poultry species but are
notable due to their zoonotic potential. Macrorhabdosis
in quail and other avian species is caused by infection
of the proventriculus and ventriculus with the yeast,
Macrorhabdus ornithogaster. Clinical signs include emacia-
tion, prostration, anorexia, cachexia, and death; whereas
histopathologic lesions are marked lymphoplastic and
heterophilic inflammation of the proventriculus and
ventriculus.
5. Parasites
Like other gallinaceous birds, Japanese quail can host a
number of internal and external parasites. Due to the lim-
ited scope of this chapter, readers are referred to a more
complete review of these topics (Reed and Jack, 2013).
1104 22.  Japanese Quail as a Laboratory Animal Model
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LABORATORY ANIMAL MEDICINE

LABORATORY
ANIMAL
MEDICINE
THIRD EDITION

Edited by

James G. Fox (Editor-in-Chief)

Division of Comparative Medicine, Massachusetts Institute of Technology, Cambridge, MA, USA

Lynn C. Anderson
Global Animal and Comparative Medicine, Covance Laboratories Inc., Madison, WI, USA

Glen M. Otto
Animal Resources Center, University of Texas at Austin, Austin, TX, USA

Kathleen R. Pritchett-Corning
Harvard University, Faculty of Arts and Sciences, and
Department of Comparative Medicine, University of Washington, Cambridge, MA, USA

Mark T. Whary
Division of Comparative Medicine, Massachusetts Institute of Technology, Cambridge, MA, USA

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