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236  |  Yao and Nair

          tolerance to the virus and after hatching make up the V+A– class,   myeloblasts comprising up to 75% of peripheral blood cells and
          with high levels of virus in the blood and tissues and an absence of   forming a thick buffy coat and usually an anaemia and thrombo-
          antibodies. By 22 weeks of age, up to 25% of meat-type chickens   cytopenia.
          exposed to ALV-J at hatch were found to be V+A–, although this   Tumours of myelocytomatosis (myelocytic myeloid leuko-
          could be affected by a number of factors including the virus strain   sis) are distinctive and can be recognized on gross examination
          (Pandiri et al., 2007).                               with some degree of certainty. Characteristically, they occur on
                                                                the surface of bones in association with the periosteum and near
                                                                cartilage, although any tissue or organ can be affected. Myelocy-
          Spread of infection                                   tomas often develop at the costochondral junctions of the ribs, on
          Birds from infected flocks, and products, such as eggs and meat,   the inner sternum, pelvis, and on the cartilaginous bones of the
          could spread infection to other flocks and locations. Because ALV   mandible and nares. Flat bones of the skull are also commonly
          is vertically transmitted, infection can spread transnationally in   affected. Tumours may also be seen in the oral cavity, trachea,
          hatching eggs and day-old chicks, and potentially through the   and in and around the eye (Pope et al., 1999). The tumours are
          semen. Importers need to guard against the introduction of ALV   usually nodular and multiple, with a soft, friable consistency and
          by requiring health certificates from the exporter relating to the   of creamy colour. In the disease caused by subgroup J ALV, myelo-
          specific disease and infection status of source flocks. Similarly,   cytomatous infiltration often causes enlargement of the liver and
          there has to be a guard against the introduction of ALV through   spleen and other organs, in addition to skeletal tumours (Wil-
          live cell culture vaccines. Introduction of a new ALV into sus-  liams et al., 2004) and myelocytic leukaemia (Payne et al., 1991).
          ceptible poultry populations could have serious consequences   Haemangioma is found in the skin or in visceral organs in
          because of absence of immunity and of effective control measures   chickens of various ages. They appear as blood-filled cystic masses
          in breeding and production populations other than at the primary   (blood blisters) or more solid tumours and consist of distended
          breeding level.                                       blood-filled spaces lined by endothelium or as more cellular,
                                                                proliferative, lesions (Campbell, 1969). They are often multiple
                                                                and may rupture, causing fatal haemorrhage (Soffer et al., 1990).
          Clinical features and pathogenesis                    More recently, many workers have reported the incidence hae-
          Outward signs of the leukotic diseases are mostly non-specific.   mangiomas in layer chickens infected with ALV-J in China (Lai
          They include loss of appetite, weakness, diarrhoea, dehydration,   et al., 2011; Pan et al., 2011; Zhang et al., 2011). Renal tumours
          and emaciation. In LL especially, there may be abdominal enlarge-  may cause paralysis due to pressure on the sciatic nerve. Sarco-
          ment. The comb may be pale, shrivelled, or occasionally cyanotic.   mas and other connective tissue tumours may be seen in the skin
          In erythroblastosis and myeloblastosis, haemorrhage from feather   and musculature. When advanced, these various other tumours
          follicles also may occur. After clinical signs develop, the course is   may be accompanied by the non-specific signs given previously.
          usually rapid, and birds die within a few weeks. Other affected   Benign tumours may follow a long course, malignant tumours a
          birds may die without showing obvious signs.          rapid one.
            Fully developed LL occurs in chickens of about 4 months of   In osteopetrosis, the long bones of the limbs are commonly
          age and older. Grossly visible tumours almost invariably involve   affected. Uniform or irregular thickening of the diaphyseal or
          organs  such  as  the  liver,  spleen,  and  bursa  of  Fabricius.  Other   metaphyseal regions can be detected by inspection or palpation.
          organs including kidney, lung, gonad, heart, bone marrow, and   The affected areas are often unusually warm. Birds with advanced
          mesentery also may show lesions. The tumour cells are B-cells,   lesions have characteristic ‘bootlike’ shanks. Affected birds usu-
          expressing immunoglobulin M (IgM) and other B-cell mark-  ally are pale, stunted and walk with a stilted gait or limp. In recent
          ers, which originate in the bursa of Fabricius but subsequently   years, ALV has also been shown to be associated with the ‘so
          develop metastasis to other visceral organs.          called fowl glioma’ (Ochiai et al., 1999), associated with cerebel-
            Natural cases of erythroblastosis (erythroid leukosis) usually   lar hypoplasia and myocarditis (Iwata et al., 2002; Hatai et al.,
          occur in birds between 3 and 6 months of age. The liver and kidney   2005, 2008; Toyoda et al., 2006; Nakamura et al., 2011).
          are moderately swollen, and the spleen often is greatly enlarged.
          The enlarged organs are usually cherry red to dark mahogany and
          are soft and friable. The bone marrow is bright red and liquid.   Immune responses
          Affected birds are often anaemic, with muscle haemorrhages and
          occasionally abdominal haemorrhage from a ruptured liver. The   Active immunity
          disease is an intravascular erythroblastic leukaemia.  Immune responses to oncogenic viruses including ALV have
            Myeloblastosis  (myeloblastic  myeloid  leukosis)  is  predomi-  been reviewed (Nair, 2013). Under natural conditions, most
          nantly a sporadic disease of adult chickens. The liver is greatly   chicks become infected by exogenous ALV from penmates or
          enlarged and firm with diffuse greyish tumour infiltrates, which   their surroundings and, after a transient viraemia, develop virus-
          give a mottled or granular (‘Morocco leather’) appearance. The   neutralizing antibodies directed against virus envelope antigens
          spleen and kidneys are also diffusely infiltrated and moderately   that rise to a high titre and persist throughout the life of the bird.
          enlarged. The bone marrow is replaced by a solid, yellowish-  The virus-neutralizing antibodies serve to restrict the amount
          grey  tumour  cell  infiltration.  A  severe  leukaemia  exists,  with   of virus in the bird, which in turn, will limit neoplasia, but they
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