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An Overview of Infectious Bronchitis Virus in Chickens
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Abbreviated title: Infectious bronchitis virus in chickens
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Summary
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Infectious bronchitis virus (IBV) is one of the foremost causes of economic loss within the
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poultry industry. IBV is a commonly occurring, economically significant pathogen of
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commercial chicken. Economic consequences to the poultry industry comprise mortality,
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growth retardation and high condemnation rates in meat-type birds. In addition, decreased egg
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production, reduced internal and external egg quality, and reduced hatchability have been
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documented in layers and breeders affecting the performance of both meat-type and egg-
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laying birds. Apart from this some nephropathogenic strains cause kidney damage. Secondary
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pathogens can complicate the disease resulting in increased morbidity and mortality. Being a
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single stranded RNA virus, IBV has an enormous capacity to change both by spontaneous
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mutation and by genetic recombination resulting into the emergence of new variants. Since
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after the first isolation of virus in 1937, it is found in almost all over the world. In addition,
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most countries are now known to have their own indigenous IBV variants. Despite the use of
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currently available live and inactivated vaccines, one of the most important difficulties to
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control IB is related to emergence of variant strains.
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Key words: Infectious bronchitis virus, chicken
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Introduction
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Infectious bronchitis virus (IBV) is an acute and highly contagious viral disease causing
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severe economic losses for those involved in the chicken industry (Cavanagh and Gelb,
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2008). IBV was initially found to cause only respiratory disease but over time the virus has
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been isolated from various non-respiratory tissues including kidneys and different parts of the
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oviduct and alimentary tract (Alexander and Gough, 1977; Ambali and Jones, 1990; Benyeda
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et al., 2009; El‐Houadfi et al., 1986; Ganapathy et al., 2012; Maiti et al., 1985; Wang et al.,
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1997; Yu et al., 2001). To date, the virus only tends to cause disease in chickens even though
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IBV-like coronaviruses have been isolated from non-chicken birds including some migratory
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birds (Cavanagh, 2007; de Wit et al., 2011a; Felippe et al., 2010). All age groups, male and
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females are susceptible to IBV (Cook and Mockett, 1995). The occurrence and severity of the
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disease is dependent on factors related to the virus, host and environment. In spite of constant
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efforts to control the disease by use of live and inactivated vaccines, the virus continues to
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cause high production losses, poor health and raises welfare concerns. In most countries,
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irrespective of chicken types, all are vaccinated using live and inactivated IBV vaccines.
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The outbreaks of infectious bronchitis are frequent due to the continual emergence of variants.
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This review summarizes in general the disease, different virus strains, its replication and
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transmission, the disease incidence and control measures for infectious bronchitis virus
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infection.
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The Disease
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In IBV infected flocks the morbidity rate can reach 100% but the mortality rate depends on
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the presence of secondary infection, flock age, immune status, management and
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environmental factors. In young chickens the mortality rate is typically 25-30% but it can
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approach 80% as it depends on the virulence of the strain. Even though all age group of
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chicks are susceptible to IBV, baby chicks are more susceptible than older ones (Cavanagh
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and Gelb, 2008).
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The most notable symptoms are those which affect the respiratory tract hence named as
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infectious bronchitis. The severities of the clinical signs are influenced by several factors
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associated with the field virus (such as strain, virulence and dosage), the host (age, sex, type
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and immune status), the environment (dust, ammonia and stress) and management and
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biosecurity levels (Ganapathy, 2009). The other forms of the disease are mainly associated
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with the reproductive, renal and gastro-intestinal systems. It has been reported that the virus
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can cause infertility in male chickens (Jones, 2010a). The virus has been isolated from the
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testicles and semen of infected males birds (Gallardo et al., 2011).
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The typical signs of IBV in chickens of less than six weeks of age are depression, huddling
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under the heat source, difficulty with breathing (often more noticed at night when the birds
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are resting), gasping, coughing, tracheal râles, nasal discharge, lethargy, watery eyes and
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mild swollen sinuses in severe conditions (Cavanagh and Gelb, 2008; Ganapathy, 2009).
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Baby chicks may die due to IBV and in uncomplicated infections chicken more than five
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weeks of age the disease may not be serious. In any age groups if IBV is mixed with other
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pathogens more severe diseases can occur (Cavanagh, 2007).
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Following infection via the respiratory route the virus spreads to other tissues including the
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reproductive tract. Infection at an early age may cause permanent damage to the reproductive
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tract leading to a decrease in quantity and quality of eggs (Benyeda et al., 2009; Ganapathy,
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2009; Raj and Jones, 1997). Infection of laying birds, results in decreases in egg production
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and the extent of the damage can vary depending of the time of the infection and chicken’s
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health status (Cook and Mockett, 1995). Affected flocks are unlikely to return to the normal
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level of production (Benyeda et al., 2009; Ganapathy, 2009).
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The nephritic form of IBV mostly occurs in broiler chickens but it might also affect young
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growing pullets and even layers. Litter can be very watery and may contain excessive urate.
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After a few days of infection the droppings of infected birds become completely wet and birds
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may die suddenly within the first ten days (Meulemans and van den Berg, 1998). Infection
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with IBV was believed to cause deep pectoral myopathy as another atypical clinical finding in
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chickens (Raj and Jones, 1996). IBV serotype 793B was isolated from a broiler breeder flock
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which had a bilateral myopathy affecting both deep and superficial pectoral muscles. It is
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characterised by paleness and swelling of the deep pectoral muscles with the presence of
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occasional facial haemorrhages (Gough et al., 1992; Raj and Jones, 1996, 1997).
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Droual and Woolcock (1994) reported that a commercial broiler flock in the Central Valley of
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California experienced respiratory disease and signs of swollen head syndrome (SHS). IBV
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M41 was isolated from the tissue samples as well as serum samples was positive for IBV but
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negative for avian metapneumovirus (aMPV). This finding suggests that SHS can also be
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associated with IBV other than aMPV (Droual and Woolcock, 1994). In an experimental
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infection with IS/885/00 in commercial broilers, head swelling was found at 15 days post-
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infection (dpi). Following necropsy, sterile incision of skin was made, and swabs were taken
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from tissues underneath the skin, and this was positive for the IS/885/00 by RT-PCR and
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virus isolation (Awad and Ganapathy, unpublished).
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Virus replication and spread
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IBV has an incubation period of 24 to 48 hours and viral spread occurs rapidly among
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chickens in a flock by aerosol and mechanical means. The initial replication of IBV occurs in
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the upper respiratory, causing the loss of ciliated epithelial cells of sinuses and trachea. In
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addition, some IBV strains also replicate in tissues along the alimentary tract (esophagus,
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proventriculus, duodenum, jejunum, bursa of Fabricius, caecal tonsils, rectum, and cloaca)
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without significant pathological effects (Boltz et al., 2004; Raj and Jones, 1997; Villarreal et
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al., 2010).
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Renal damage associated with different IBV strains is an increasingly important feature of
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IBV infection especially in broilers as nephropathogenic strains produce less respiratory signs
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(Ziegler et al., 2002) and lesions (Glahn et al., 1989), but can induce high mortality (de Wit et
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al., 2011b). Nephropathogenic IBV outbreaks can be complicated and/or exacerbated by
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management problems such as high dietary calcium (Glahn et al., 1989). Infection of enteric
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tissues does usually not manifest itself clinically but persists for long periods and results in
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fecal virus excretion. It has also been reported that the virus replicated in the gut for longer
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than in the respiratory tract (Jones, 2010b). In recent years some strains of IBV have caused
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proventriculitis (Benyeda et al., 2010; de Wit et al., 2011a; Gough et al., 2008; Sun et al.,
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2011; Yu et al., 2001) and enteritis (Jones, 2010a). Field infection with 793B is associated
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with enteritis (Jones, 2010a). Apart from this, IBV replicates in the oviduct and testes of
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infected chickens leading to decreases in egg production and fertility (Boltz et al., 2004;
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Grgic et al., 2008; Villarreal et al., 2007). In breeder and laying hens the arrival of IBV QX
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has reinforced the ability of some IBV strains in causing substantial pathological damage to
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female chicken’s reproductive tracts. This in turn causes delayed onset of production, poor
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peak in egg production, high percentage of false-layers and poor quality of eggs (de Wit et al.,
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2011b; Ganapathy, 2009).
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Infectious bronchitis virus and strain variations
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IBV is caused by a coronavirus, of the family Coronaviridae and over twenty five known
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genotypes worldwide (Snyder, 2002). It is an envelope and single stranded RNA virus. Three
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virus-specific proteins have been recognised: (i) the spike (S) glycoprotein; (ii) the membrane
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or matrix (M) glycoprotein; and (iii) the nucleocapsid (N) protein (Lai and Cavanagh, 1997).
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Being a single stranded RNA virus, IBV has an enormous capacity to change both by
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spontaneous mutation and by genetic recombination (Cavanagh and Gelb, 2008) . When these
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events occur with IBV both are most likely to result in the emergence of new variants if they
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occur in the hypervariable regions of the spike gene. Most countries are now known to have
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their own indigenous IBV variants which are characterised and are frequently named from the
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laboratory or researcher in which or by whom they were first isolated for instance
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Massachusetts, Connecticut, Arkansas, D274 and QX. It is very well understood that whilst
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many new variants are unable to replicate or survive for long time, a few manage to emerge
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and become of economic importance either worldwide or in restricted geographic areas.
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Host range
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It is accepted that chickens are the most significant natural hosts of avian infectious
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bronchitis. However, farmed pheasants have also been found to be natural hosts for IBV
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(Ignjatovic and Sapats, 2000). With history of clinical respiratory disease and reduce egg
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production, this suggests that pheasants could be second natural hosts for IBV(Cook and
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Mockett, 1995). However, it appears that some species of pheasant are not susceptible to the
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virus or possibly some strains of IBV could infect pheasants (Ignjatovic and Sapats, 2000).
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Recently, there is more evidence which suggests that IBV has a wider host range than was
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previously thought and is not only limited to galliform (chicken-like) birds(Cavanagh, 2007;
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Cavanagh and Gelb, 2008), but other species such as geese, ducks, and pigeons might also
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play a role in the spread of IBV strains over the world (de Wit et al., 2011a; Felippe et al.,
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2010) . Interestingly the role of the wild birds in the world of IBV is largely unknown and
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speculative but it is certainly a critical area that deserves more attention and research in near
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future.
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Incidence and geographical distribution
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In 1931 Schalk and Hawn reported the first occurrence of IBV in the United State, as a "new
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respiratory disease of baby chicks" (Schalk and Hawin, 1937). The first isolate of IBV was
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the Beaudette strain (Beaudette and Hudson., 1937). Later, the M4 1 strain was isolated in the
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North Dakota and found to be serologically related to Beaudette strain (Bracewell, 1975).
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Strains of this serotype were among the first to be made into live vaccines, e.g. H 120
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(derived from a Dutch isolate of 1955) (Bijlenga et al., 2004).
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The Australian 'T' strain was the first IBV strain isolated in Australia in 1962 (Cumming,
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1963; Ignjatovic et al., 1997). Many different IBV variants have been isolated and
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characterized since then and IBV has always evolved independently here from the rest of the
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world due to its geographical isolation (Ignjatovic et al., 2006).
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In Europe, only 30% of the isolates were serologically related to the known North American
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serotypes in the 1970s. The majority of the remainder strains were found to be related to four
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Dutch strains; D207, D212, D3128 and D3896 (Davelaar et al., 1984), by VN test. Many of
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the British IBV strains isolated between 1981 to 1983 were closely related to the Dutch
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Strains; D207, 03128 and D3896 (Cook, 1984). Although these viruses caused respiratory
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disease and aberrant drop in egg production in layer flocks, but, they were not associated with
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poor egg quality (Cook, 1984; Cook and Huggins, 1986). Thereafter, many IBV variants were
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isolated, from the UK (Gough et al., 1996), and in other European countries including France
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(Picault et al., 1986) Belgium (Meulemans et al., 1987), Italy (Capua et al., 1994; Zanella et
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al., 2000; Zanella et al., 2003) Poland (Minta et al., 1998) and Spain (Dolz et al., 2006,
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2008). Among these of major importance internationally was the variant called variously
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793B that emerged in the1990s and quickly spread to many parts of the world (Gough et al.,
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1992; Parsons et al., 1992). In Brazil, in addition to the local strains, viruses similar to the
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European 793B type, Mass-type and Arkansas viruses have been reported (Villarreal et al.,
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2010).
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In 2002 sequence of a new IBV genotype named Italy 02 with economical importance was
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submitted to the GenBank nucleotide database. By using virus neutralization assays it was
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demonstrated
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reference IBV serotypes, and indicated that Italy 02 is likely to be a new serotype (Dolz et al.,
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2006). Later on, Italy 02 has been described to be the third most frequently detected IBV
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strain and probably the dominant wild type in Western Europe. The origin of Italy 02 is
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unknown, but it has been spreading predominantly throughout Europe (Worthington et al.,
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2008).
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By the mid-1990s many published reports have revealed the diversity of IBVs causing disease
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in different part of China. But, possibly the most significant IBV variant emerged from China
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is the QX variant reported in association with proventriculitis (Wang et al., 1998). IBV QX
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later appears to have spread to Russia and then to Europe (Bochkov et al., 2006; Worthington
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et al., 2008). This genotype of IBV mainly affects the respiratory and reproductive systems
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(Ducatez et al., 2009; Terregino et al., 2008). IBV QX isolated in Netherlands also causes
that little antigenic relatedness exists between Italy 02 and some of the
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renal disease and false layers (Irvine et al., 2010; Terregino et al., 2008). In the UK, IBV QX-
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like was isolated from proventriculus of broilers with poor feed conversion ratio (FCR) and
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body weight gain also showing respiratory signs (Ganapathy et al., 2012).
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Another IBV genotype, the Q1, which was genetically and serologically distinct from IBV
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classical strains, was reported in China from 1996 to 1998 (Yu et al., 2001). Recently, Q1
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IBV strain has
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(Jackwood, 2012), Middle East and Indian subcontinent (Ganapathy, unpublished)
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Of late, IBV genotypes with certain regional importance such as IBV IS/855/00 (Meir et al.,
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2004), and IS/1494/06 were isolated in Israel (Meir, R. Personal communication). Later on
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these strains were also reported in the Turkey (Kahya et al., 2013), Egypt (Abdel-Moneim et
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al., 2012), Iraq (Mahmood et al., 2011) and Libya (Awad and Ganapathy, unpublished).
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In nutshell, variants of most importance in major parts of the world, such as 793B or QX that
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have spread over Asia, Europe and Africa in a short period have not been reported in the USA
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or Australia. On the other hand, the major strain of the USA, the Arkansas strain has hardly
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been reported outside the USA. A pathogenic strain like D1466 that has been in some
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countries in Western Europe for three decades now has hardly been reported outside Western
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Europe, although it is very difficult to achieve a sufficient level of protection (Cook et al.,
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1999), against this strain, which would make it easier for the strain to spread to other areas. It
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seems likely that geographical isolation and control measures employed in countries may play
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a part in preventing entry of IBV variants (de Wit et al., 2011a).
also been identified in Europe (Toffan et al., 2011), South America
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Prevention and Control
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Since the discovery of the disease, past and present control strategies have struggled to
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control the disease due to existence of numerous IBV strains more of which continue to
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emerge. With such diversity it is becoming increasingly difficult to control the infection
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using the currently available live and inactivated vaccines. Other than biosecurity and
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management improvements these vaccines are in use worldwide. There is a continuous
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emergence of variants through natural recombination. Even though most of these variants
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disappear, few tend to stay and continue to cause disease resulting in production loss (de Wit
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et al., 2011a). This appears to be due to an inability of currently available vaccines or
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vaccination programmes in tendering full protection against emerging strains. In recent years,
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IBV QX has become a huge threat to the poultry industry despite high and proper use of IBV
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vaccine and vaccination in the Western Europe. It appears that current vaccines and
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vaccination strategies have managed to reduce the impact of these newly emerged IBVs, in
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terms of the occurrence of disease and production losses but has allowed the continuous
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spread of the viruses.
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The appropriate strategy for control of IBV is to use vaccine strains that are similar to those
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found in a particular farm or area. In areas where this is not possible or where there were no
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vaccines available for the prevalent strains, vaccination programmes consisting of different
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strains tend to induce a higher and broader protection against challenges with several
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heterologous strains (Cook et al., 2001; Cook et al., 1999; Terregino et al., 2008). Generally,
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the success rate would depend on the genetic differences between the vaccine viruses and the
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field viruses. Live vaccines are mostly administered by spray or drinking water (Cavanagh
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and Gelb, 2008), but for better flock vaccine take it is preferable to vaccinate intraocularly or
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intranasally. In long-term birds, for example breeders and layers, and in some countries in
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broilers, IBV inactivated vaccines are used. This vaccine is given by intra-muscular or
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subcutaneous injections for induction of high levels of humoral antibodies, as would protect
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the birds against invading IBVs and avoids drops in egg production or quality. Such
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vaccination in breeder birds helps in ensuring transfer of high and uniform levels of maternal
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antibodies to hatching chicks. Maternal antibodies are important in conferring some
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protection against of virulent IBVs (Davelaar and Kouwenhoven, 1977; Mondal and Naqi,
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2001).
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Conclusion
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Avian infectious bronchitis still poses a great challenge for the chicken industry worldwide.
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As a virus that goes through continuous changes, increasing number of regional and global
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variants has been identified. But the foregoing survey of available knowledge does not ask for
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new vaccines at every stage. Fortunately some of the live vaccines give broad protection
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against most of the variants reported to be causing respiratory/ nephrotropic IB. There is a
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promising choice of incorporating strategic strains in inactivated vaccines. Apart from this, it
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appears that continuous active surveillances, improved management and bio security, are all
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essential to suppress losses due to IBV infections.
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