Domestication changes in Japanese quail (Coturnix japonica)
The
ancestor of domesticated quail is the wild Japanese quail (Coturnix japonica),
which belongs to genus Coturnix, family Phasianidae, order Galliformes, class
Aves (Chang et al. 2007). Quail have been reared from ancient times in East and
South East Asia as fighting, song and decorative birds. The earliest available
records of using quail as song birds comes from China, dating back to 770-476 BC
(Chang et al. 2005). In Japan, quail were reared from 12th century for their
song (Kovach 1974). Japanese quail eggs and meat were used as a food in Asia
from the 17th century onward (Genchev 2014). At the end of 19th and beginning of
20th century, the true domestication of Japanese quail occurred, leading to
serious changes in egg productivity (Howes 1964; Donchev and Angelov 1971; Mills
et al. 1997). Later, in the second half of the 20th century, meat type quail
were selected in North America and Western Europe, when the quail farming became
industrial (Genchev 2014). Nowadays, quail can be found in almost any part of
the world, where they are used as a productive, ornamental, fighting, singing or
laboratory animals (Mills et al. 1997; Minvielle 2004; Pavlova et al. 2018;
Lukanov et al. 2018a). In order to avoid confusion of names for the wild and
domestic forms, Lukanov (2019) proposed the term ‘domestic quail’ and a
corresponding Latin name C. j. domestica, as a clear differentiation from its
wild ancestors (C. japonica).
Due to selective breeding, there are some major differences between their wild
ancestors, mostly visible in morphological, behavioural and productivity traits
in domes¬tic quail. These reflect greater intraspecific differences, even more
so than those between species in the genus Coturnix.
The aim of this review was to emphasise the changes that occurred during the
domestication of Japanese quail, supporting the proposal of using the term
domestic quail (C. j. domestica).
Domestication-induced changes
Domestic quail have been widely used as productive, experimental or ornamental
animals for more than a century, and much longer as singing or fighting birds.
The multifunctionality of this small bird has transformed it in different
directions, which moves it further away from its ancestor.
Morphological changes
Plumage colour, feather growth and structure. One of the first
morphological changes in domesticated quail was associated with plumage colour.
A number of mutations in plumage of domestic quail have been described. These
affect the primary (background) or secondary (the feather pattern) colour. Among
the commonest colours are variants of golden (Y), tuxedo (wb, cr, wp), extended
brown (E) and recessive white (wh). Other plumage colours have been reported in
this species, such as dominant black (Cheng and Kimura 1990), recessive black (Hiragaki
et al. 2008), redhead (Truax and Siegel 1981), pancy (Tsudzuki and Wakasugi
1987), fawn-2 (Tsudzuki 1996) and lavender/blue (Cheng and Kimura 1990;
Minvielle et al. 2002). Some plumage patterns are associated with lower
liveability, such as black at hatch (Minezawa and Wakasugi 1977; Shiojiri et al.
1999; Niwa et al. 2003), buff (Sittmann et al. 1966), white (Wakasugi and Kondo
1973), marbled plumage (Yakovlev et al. 1975) and orange mutation (Ito and
Tsudzuki 1994). Sex-linked colour mutations have the possible application in
autosexing quail, including imperfect albino (Cheng and Kimura 1990), two brown
varieties (roux and red (Minvielle et al. 1999, 2000)) and lavender plumage (Fulton
et al. 1982; Gunnarsson et al. 2007; Lukanov and Genchev 2017; Lukanov et al.
2018b). Other colour mutations identified in Japanese quail have been described
in detail by Cheng and Kimura (1990), Mizutani (2003) and Tsudzuki (2008).
A total of nine mutations affecting feather growth and structure are
acknowledged in Japanese quail (Tsudzuki 2008). Six of them are determined by
autosomal recessive genes for ruffle (rf), porcupine (pc), short barb (sb),
downless (dl-1 and dl-2), curly (CU*C) and defective feathers (Df and mdf). Two
mutations are due both to a recessive and dominant autosomal gene, which are
defective feathers (Df, mdf) and fray (Fr, mod). Only one, the partial
featherlessness (Pf), is due to an autosomal dominant factor. Other plumage
mutations reported in domesticated Japanese quail are associated with the
presence of an ear or throat tuft, as summarised by Cheng and Kimura (1990) and
Tsudzuki (2008).
Live body weight and size. Considerable changes have occurred in live
weight and body size. Average live weight of wild Japanese quail varies within
85-110 g (Wilson et al. 1971; Chang et al. 2009; McGowan and Kirwan, 2020).
Normally, males are lighter than sexually mature females (Mills et al. 1997;
Chang et al. 2005). This live weight difference is not reflected in body
proportions, such as wingspan and body length (Johnsgard 1988). Domestic quail
are bigger than wild Japanese quail (Mills et al. 1997), and have large
variation in body weight in relation to the productive type. Thus, three
productive types can be distinguished; light (egg-laying), heavy (meat-type) and
dual-purpose. Quail reared for the production of hatching eggs are similar to
the original Japanese type domestic quail, with a live weight slightly higher
than wild breeds. According to Mizutani (2003), breeding quail in Japan weigh
100-130 g (males) and 120-160 g (females). Chang et al. (2009) reported average
live weights of domestic quail in China to be 106.8 ± 26.6 g for males and 134.3
± 14.3 g in females, which is about 45% more than in their ancestors. At the
other extreme are quail for the production of meat, whereby body weight is more
than 250% higher than that of wild breeds, being about or over 300 g (Minvielle
2004; Tavaniello 2014). Dual-purpose quail are predominantly raised by hobby
breeders, and in Europe and Russia for egg production. Their live weight is
about 200-250 g, which is representative of the Estonian breed (Lember and Laan
2013).
Other morphological changes. A study on phenotypic traits of wild and
domestic Japanese quail showed differences in body shape, tarsus length, tarsus
girth, wing length and body slant length (Wilson et al. 1971; Chang et al. 2001,
2009). The authors did not report any differences in plumage pattern or the size
and shape of the beak.
Differences between wild and domesticated quail are found in digestive tract
devel¬opment. Mihaylov and Dimitrov (2010) reported higher relative weight of
the intestinal tract in domestic Japanese quail as compared to wild and hybrid
forms. The duodenum of wild and hybrid quail was longer in comparison to
domesticated breeds.
Behavioural changes
Investigation into behavioural traits of wild quail is difficult, due to the
nature of their biology and habitat (Bannerman 1963; Krause 2003; Sanchez-Donoso
et al. 2018). Whereas, domesticated breeds are a far more convenient subject for
ethological studies (Farris 1964; Wilson and Bermant 1972; Schmid and Wechsler
1997; Galef et al., 2006; Labaque et al. 2008; Ball and Balthazart 2010; Santos
et al. 2017).
Migratory behaviour. Domesticated Japanese quail do not exhibit migratory
behaviour, unlike their wild ancestors and common quail. These changes due to
domestication are marked and observed even in hybrids (Deregnaucourt 2000;
Deregnaucourt et al. 2005a; Huisman 2006). Deregnaucourt et al. (2005b) observed
migratory tendencies in two populations of domesticated Japanese quail, which
was attributed to the introduction of European quail (Coturnix coturnix)
genetics. During the 1970s, the introduction of ‘wild blood’ (i.e. genetics) in
domestic quail used for hunting was a popular practice aimed at improving flying
along with preserving the higher body weight and size of birds (Nadal 1992;
Mills et al. 1997; Deregnaucourt et al. 2005b).
Sexual behaviour. Sexual behaviour in wild Japanese quail in their
natural environment is relatively poorly studied, as experiments in an
artificial environment usually fail (Nichols 1991). Quail are considered
monogamous (Schwartz and Schwartz 1949; Stevens 1961; Kawahara 1967) or socially
monogamous birds (Nichols 1991), but some authors consider them polygamous (Dementiev
et al. 1952; Domjan and Hall 1986). A review by Kovach (1974) concluded that the
mating of wild Japanese quail is in a transitional state between polygamy and
monogamy. In the view of Mills et al. (1997) Coturnix japonica may show regional
variation in mating patterns. In the field, the breeding period of quail is
during the spring (Pappas 2002), when increased duration of daylight stimulates
the growth of gonads and sex hormone concentrations (Sharp 1984). Other
environmental factors promoting mating include the elevation of ambient
temperature and access to food (Nichols 1991). These physiological changes are
associated with the behaviour of wild quail, although, unlike them, domestic
breeds do not exhibit a seasonal pattern in reproduction, due to controlled
microclimatic parameters and photostimulation opportunities (Mills et al. 1997).
Placed under natural conditions, the sexual function in domestic quail declines
during the winter due to low ambient temperatures and short duration of daylight.
Even though domestic quail are polygamous birds, the recommended sex ratio for
rearing is 1:2 to 1:4, depending on the production type (Cheng et al. 2010;
Genchev 2014), and monogamy has been reported (Orcutt and Orcutt 1976; Nichols
1991). Ottinger and Brinkley (1979) reported first mating attempts in domestic
quail at 35 days of age and true copulation: after 37 days of age. In an earlier
report, the peak in copulation occurred at 52 days of age (Ottinger and Brinkley
1978). Stefton and Siegel (1973) demonstrated that the highest sexual activity
in male domestic quail was exhibited between 70 and 210 days of age. As
productive age advances, the libido of males declines, but not the sexual
activity of females at the same age (Woodward and Abplanalp 1967; Ottinger and
Balthazart 1986). Domestic male quail and F1 crosses had a considerably higher
libido that wild males (Nichols 1991; Chang et al. 2009). The peak sexual
activity occurred in early afternoon in both domestic and F1 male quail, and
before noon in wild breeds. Ottinger et al. (1982) observed peak sexual activity
in the early afternoon and the lowest during the late afternoon, when plasma
testosterone concentrations were lowest (Ottinger and Brinkley 1978). Comparably
to these reports, Nichols (1991) reported higher levels of mating behaviour
traits in domestic compared to wild quail.
Nesting and brooding behaviour. Wild quail exhibit nesting and brooding
behaviour from April-May to August, e.g. the natural breeding period (Pappas
2002). Conversely, nesting and brooding instincts are absent in domestic quail,
and only sporadic reports have documented nesting and brooding instincts (Stevens
1961; Orcutt and Orcutt 1976; Nichols 1991). Through hormonal treatment with
progesterone, McCollam and Schein (1974, as quoted in Orcutt and Orcutt 1976)
succeeded in provoking nesting and brooding in domestic Japanese quail. Hormonal
treatment with prolactin, combined with oestradiol or testos¬terone, promoted
brood patch development, but not brooding behaviour (Hohn 1981). Brody (1970, as
quoted in Mills et al. 1997) did not succeed in encouraging nesting and brooding
instincts in female quail treated with oestradiol, progesterone, prolactin and
luteinising hormone, both independently or simultaneously. Hybrids of wild and
domes¬tic quail exhibit nesting and brooding behaviour, but these are unstable
compared to those of the wild form, therefore their nests were exposed to a
greater risk of predator attacks (Capdevila et al. 2016). According to Nichols
(1991) the parent-offspring rela¬tionship breakdown occurs earlier in domestic
than in wild quail. The author demon¬strated that, in the field, domestic quail
were significantly more quiet than wild birds, which were more timid and
vigilant. There is evidence that fear responses in quail are genetically
determined (Mills et al. 1997) and there have been reports for successful
selection of that trait (Kiker et al. 1976; Bessei 1979).
Other behavioural changes. Other reactions in Japanese quail are
influenced by domestication, including vocalisa¬tion, mating calls, aggression
and fighting. Vocalisation differs between C. coturnix and C. japonica (Guyomarc’h
and Guyomarc’h 1996; Collins and Goldsmith 1998; Deregnaucourt and Guyomarc’h
2003; Deregnaucourt et al. 2005b; Huisman 2006). According to Nichols (1991),
the sounds made by domestic and wild male Japanese quail were different, while
according to Chang et al. (2009) they were identical. Deregnaucourt et al.
(2009) detected high inter-individual variations in sounds emitted by male
domestic quail. Chang et al. (2009) found that, out of the breeding season,
vocalisation in wild Japanese quail occurred mainly at the time of feeding.
Specific sounds could be heard when birds were frightened, and during the
breeding season, males emit frequently specific mating calls, which are
different for paired and unpaired males. Vocal activity was about 30 times more
intensive in domestic Japanese quail compared to wild breeds. This was in line
with results reported by Nichols (1991).
A similar relationship was discovered with regard to aggression and fighting,
with F1 crosses being closer to domestic quail behaviour (Chang et al. 2009).
Nichols (1991) did not find any differences in aggression between wild and
domestic quail, and observed higher activity in males of both types during the
egg production cycle of females. Male wild quail exhibited higher aggression
towards females than domestic breeds at the time of pair formation. Aggression
was most commonly territorial and mate-guarding, and male birds were more active
due to higher testosterone levels (Wingfield et al. 1990; Trainor et al. 2004).
The aggression in quail could be pathological, and has been associated with
cannibalism and pterygophagy episodes (Pizzolante et al. 2006; Cheng et al.
2010). This behaviour is multifactorial, due to genetic, environmental factors,
erratic feeding and other stressors (Savory 1955; Haslam 2008).
Changes in production traits
Sexual maturity. Earlier sexual maturity is influenced by various
extrinsic and intrinsic factors. Male wild Japanese quail attain sexual maturity
at 52 days of age, and females at 63 days of age (an Age: The Animal Ageing and
Longevity Database 2017). Wilson et al. (1971) confirmed that female wild
Japanese quail enter the puberty at 59 days of age on average, but later they
mention that this happens at 117 days. According to Hoffmann (1988) young wild
quail could be considered as sexually mature at four weeks of age. Domestic
Japanese quail, placed under optimum conditions and appropriate light regimens,
attain sexual maturity at about four to five weeks of age, and females begin
regularly laying eggs at about six weeks of age (Cheng et al. 2010). Mizutani
(2003) gave an average age of sexual maturity onset of 38-42 days. From the
available literature, there are various data concerning this trait in domestic
quail, ranging from 36 to 65.5 days (Wilson et al. 1971; Sachdev and Ahuja 1986;
Thomas and Ahuja 1988; Inal et al. 1996; Drbohlav and Metodiev 1996; Gunes and
Cerit 2001; Camci et al. 2002; Ipek et al. 2003; Bahie El-deen et al. 2008;
Chimezie et al. 2017a; Elkomy et al. 2019). This variation could be explained by
the various genotypes of birds used and different feeding and rearing conditions.
Egg production. One of the main production traits, that is strongly
influenced by domestication, is egg¬laying capacity. Wild Japanese quail lay
about five to 14 eggs per clutch (Dementiev et al. 1952; Hoffmann 1988;
Johnsgard 1988), with up to three broods per year. Domestic quail can lay more
than 250 eggs per year (WIlson et al. 1961; Lucotte 1974; Mandal et al. 1994;
Lotfi et al. 2012; Genchev 2014). This major difference has been attributed
mostly to the effects of selection and, especially, the eradication of the
brooding instinct, early onset of sexual maturity, lack of seasonal reproduction
pattern and lower sensitivity to stress (Chang et al. 2009).
Egg quality traits. Another trait that has undergone major changes during
domestication of Japanese quail, is egg weight and size. Wild Japanese quail lay
eggs with weight of 7.6 g and size 29.8 x 21.5 mm (Johnsgard 1988), or,
according to Chang et al. (2009), 6.93 ± 0.80 g and size of 27.8 x 21.6 mm. The
eggs of wild Japanese quail are lighter than those of domestic breeds (Wilson et
al. 1971; Chang et al. 2009). Egg size is proportional to its weight. Depending
on the productive type, domestic quail lay eggs with average weight of 9-14 g,
which are 20 to 100% heavier than from wild breeds. Egg-laying breeds produced
smaller eggs weighing about 9-12 g (Mizutani 2003; Mori et al. 2005; Chang et al.
2009; Santos et al. 2011; Hrncar et al. 2014; Nunes et al. 2016; Chimezie et al.
2017b). The average egg weight in heavier meat-type lines and breeds ranges from
12 to 14 g (Mori et al. 2005; Santos et al. 2011; Genchev 2012; Hrncar et al.
2014; Genchev 2014; Lukanov et al. 2018a; Taha et al. 2018; Elkomy et al. 2019).
The marked progress in selection for this trait is due to the importance of egg
size for consumers and its moderate to high heritability (h2 = 0.37 ± 0.09 (Daikwo
et al. 2013), 0.59 ± 0.23 (Momoh et al. 2014), 0.83 ± 0.01 (Sezer 2007)).
According to Chang et al. (2009) eggs from domestic quail have a smoother
surface and more oval shape, while those of wild quail have an awl-like shape
and rough surface. The authors reported statistically significantly lower values
of almost all egg indices in wild quail eggs compared to those from domestic
breeds.
Some mutations affecting eggshell colour in Japanese quail are known. The normal
colour of eggshell is whitish/white with a slight brown to bluish tint,
spattered with numerous rusty brown, brown to dark brown spots of various size
and shape. Three different mutations in domestic quail are acknowledged to
influence eggshell colour, resulting in red, white and blue shells (Cheng and
Kimura 1990; Tsudzuki 2008). Two of these are due to autosomal recessive genes:
we (white eggshell) and ce (light blue eggshell, termed celadon). The third one
is autosomally dominant R (red eggshell). The colour is due to changes in the
deposition of protoporphyrin and biliverdin on the shell. Chang et al. (2009)
reported better colour patterns for eggshells from domestic than in wild quail,
the latter eggs had a dull whitish colour.
Reproductive traits. Other reproductive traits, such as fertilisation and
hatchability differ. Wilson et al. (1971) reported lower fertilisation and
hatchability in wild vs. domestic quail, reared under experimental conditions.
Comparing fertilisation rates and the hatchability of set and fertile eggs in
domestic and wild Japanese quail, Chang et al. (2009) found significantly better
reproductive performance in domestic quail. The authors reported that the
average fertility rate and hatching rate of fertilised eggs was 0.42 ± 0.10
(42%) and 0.53 ± 0.01 (53%) for wild and 0.87 ± 0.01 (87%) and 0.88 ± 0.08 (88%)
for domestic quail, respectively. Although the results for fertility and
hatchability of farmed quail eggs in the different studies vary widely, most of
them are in line with those presented by Chang et al. (2009). With respect to
studies on wild Japanese quail, some authors reported even lower values for
these reproductive parameters (Xu et al. 2003). These poor reproduction traits
in wild Japanese quail under experimental conditions could be attributed to the
relatively small number of birds used, season¬ality and stress accompanying
rearing in artificial environment (Xu et al. 2003). In contrast, Nichols (1991)
demonstrated high fertilisation of eggs laid by wild Japanese quail, reared in
big pens, at over 90%, with varied hatchability between different years, ranging
from 63% to 95%. Crossing wild males with domestic female quails and vice versa
has resulted in better fertilisation and hatchability for the former
combina¬tion (Xu et al. 2001). In fact, the differences found between wild and
domestic breeds are not due to actual changes in reproductive parameters but
rather because of behavioural characteristics of the wild quail under
experimental conditions.
Meat productivity and meat quality. The available literature reveals
fragmentary information on meat quality and growth performance in wild Japanese
quail. In contrast, there are many studies on domestic quail productivity. This
is understandable, due to the fact that the second most important quail farming
production is for meat, while bird hunting is more for sport than food. As
mentioned above, domesticated quail have bigger body weight, and so can yield
more meat than wild breeds.
Quail meat is richer in protein and leaner than most other livestock and poultry
(Prabakaran 2003). Breast meat contains 22.23-23.38% protein, with less in leg
meat at 20.49-20.91% (Genchev et al. 2008). These authors found higher fat
content in leg meat than in breast (3.26-3.39% and 2.21-2.75%, respectively).
The meat characteristics from male quail are slightly better expressed than from
females (Walita et al. 2017). The yield of meat from quail from 28 days (Lukanov
and Genchev 2018) to 35 days (Kaitazov and Genchev 2004) of age, when reaching
200-250 g live weight, is deemed profitable. The carcass yield for meat quail is
between 65% (Genchev et al. 2008; Panda and Singh 1990) and 69.5% (Tavaniello et
al. 2014), or 60-62% in skinless meat (Genchev et al. 2008, 2018). Wild quail
meat is higher in protein, Fe and Zn levels in comparison to the farmed breeds (Khalifa
et al. 2016).
Conclusions
Due to the genetic selection of domestic quail (C. j. domestica) there are major
differences between them and their wild ancestors (C. japonica). These involve
morphological, behavioural and productivity characteristics. The most
significant differences are live body weight and size, which varies
significantly within the domestic breeds due to production type (meat or eggs).
Due to selection, a number of plumage colours, other than in the wild type, have
been established and promoted due to ornamental and exhibition poultry breeding.
Domestication has had a major impact on a number of behavioural reactions, with
probably the most severely affected being nesting and brood¬ing, and the
migration response. The use of domestic quail as a productive bird has had an
impact on egg and meat production performance. A major increase has been
observed in egg production and weight. The differences in fertility and
hatchability between wild and domestic quail are mainly due to the more
difficult adaptation to the standard experimental conditions for wild birds. In
summary, the large phenotype diversity seen in domestic quail has been highly
influenced by domestication.
Disclosure statement
No potential conflict of interest was reported by the authors.
Notes on contributors
Hristo Lukanov, PhD, DVM, is a lecturer at Department of “Animal Science
- Monogastric and Other Animals”, Faculty of Agriculture, Trakia University,
Stara Zagora, Bulgaria where he has been working in Poultry Science since 2016,
mainly on domestic chickens and domestic quail. His professional interests
include Poultry meat and egg quality, Poultry genetic diversity, Poultry feeding,
etc. He published more than 80 scientific papers and other publications, and 3
books, from the field of Poultry Science and ornamental Poultry. Since 2010 he
is chairing the board of the Bulgarian Association of Poultry Breeders, and
since 2013 is a vice-president of the UFSDABB (Union of Fanciers and Small
Domestic Animal Breeders of Bulgaria).
Ivelina Pavlova, PhD, DVM, is an assistant professor at Department of
“General Livestock Breeding”, Genetics unit, Faculty of Veterinary Medicine,
Trakia University, Stara Zagora, Bulgaria. Her professional interests include
pharmacogenetics, poultry genetic diversity, poultry production, etc.
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