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1012  Section 9  Infectious Disease

            the United States, mainly the southeastern states.   The life cycle of H. americanum differs from that of H.
  VetBooks.ir  Hepatozoon canis infects the hemolymphatic tissues and   canis particularly with regard to the parasite’s target tis-
                                                              sues in the dog, the formation of distinct large cysts and
            causes anemia and lethargy whereas H. americanum pri-
            marily infects muscular tissues and induces severe
                                                              tion. Sporocysts of H. americanum sporozoites from the
            myositis and lameness [16,17]. Feline hepatozoonosis is   the inflammatory reaction formed in response to infec-
            caused by H. felis, which is prevalent in the Mediterranean   hemocoel of an infected vector tick ingested by dogs
            basin, Asia, Africa, South, Central and North America. It   release infectious sporozoites following contact with bile
            infects muscle tissues including the myocardium and has   in the gut. Sporozoites are thought to cross the gut wall,
            not been associated with severe clinical disease [18].  and then taken up by host macrophages and transmitted
            Recently, H. canis and  H. silvestris  infections have   via the lymph system or hematogenously to the target
            been described in cats in Europe.                 organs consisting of skeletal and cardiac muscles.
             In  contrast  to most  tick‐borne pathogens which  are   Hepatozoon americanum‐infected macrophages lodge
            transmitted via the tick salivary glands, Hepatozoon spp.   between striated muscle fibers in these tissues and form
            infect vertebrates when they ingest arthropod hosts con-  muscle cysts. Concentric layers of mucopolysaccharide
            taining infective sporozoites. The main vectors of  H.   material are deposited by the infected cell forming a
            canis are the ticks  Rhipicephalus sanguineus and  R.   cystic structure referred to as “onion skin cyst.” The
            turanicus worldwide and  Amblyomma ovale in South   encysted zoite undergoes merogony and multiple mero-
            America, whereas the Gulf Coast tick Amblyomma mac-  zoites are formed. Eventually, the cyst ruptures, releasing
            ulatum is the vector of H. americanum in North America.   mature individual merozoites into the surrounding tissue
            The vector of H. felis has not been identified to date but   and eliciting a strong local inflammatory response. The
            transplacental transmission in the feline host has been   pyogranulomatous inflammatory response is associated
            shown. Transplacental transmission from dam to pups   with severe musculoskeletal pain. The merozoites are
            has also been demonstrated for  H. canis infection but   taken up by leukocytes and are thought to undergo addi-
            not for  H. americanum. Transmission by carnivorism   tional merogonic cycles or enter the blood vasculature,
            and predation of the canine host on intermediate or   develop into gamonts and circulate in monocytes.
            wildlife transport hosts with ingestion of parasite cyst
            forms from their tissues has been shown for H. america-  Epidemiology
            num and is considered as a main route of dog infection
            with this parasite [19].                          Hepatozoon canis infection is reported mainly from
             The life cycle of H. canis revolves between the canine and   tropical, subtropical, and temperate climate regions
            tick hosts. When dogs ingest the vector tick or tick parts, H.   where vector tick species are abundant. Infection with
            canis sporozoites are released from sporocysts in the intes-  H. canis has also been reported in dogs in the southeast-
            tine and penetrate the gut wall. The sporozoites invade   ern United States, mainly in some regions where  H.
            mononuclear cells and disseminate hematogenously or via   americanum is also prevalent, and co‐infections of both
            the lymph to hemolymphatic target organs that include the   species have been reported. In Europe, autochthonous
            bone marrow, spleen, and lymph nodes and to other inter-  dog infection is found mostly close to the Mediterranean
            nal organs such as the liver, kidney, and lungs. Meronts in   basin. It has also been described in dogs imported into
            which asexually dividing merozoites develop are formed in   nonendemic countries such as Germany, in foxes in
            the dog’s tissues in the process of merogony and may cause   European countries where autochthonous canine infec-
            cellular necrosis in tissues. Merozoites release from mature   tion is rare or has not been reported, and in Slovakia and
            meronts, invade neutrophils and monocytes, and develop   Poland where R. sanguineus, its tick vector, is not pre-
            into gamonts. Alternatively, merozoites can produce sec-  sent. The prevalence of H. canis infection differs between
            ondary meronts in the target tissues. The tick, which serves   regions.  The  rates  of  infection  based  on  blood  smear
            as the definitive host, is infected by ingesting leukocytes   examination or PCR range from 1% to 64%.
            containing gamonts when feeding on a parasitemic dog.   The presence of H. americanum infection in the United
            Morphologically  indistinguishable  male  and  female  H.   States has expanded north and east from Texas, where it
            canis gamonts are released from the dog leukocytes within   was first detected in 1978. It is found in several southeast-
            the tick gut, associate in syzygy, and differentiate in the pro-  ern states, and occasionally cases are reported in other
            cess of gametogenesis to distinct gametes. After fertiliza-  locations including Nebraska, Vermont, Washington, and
            tion, the zygote divides and sporogony takes place with the   California. Hepatozoon americanum infects wild coyotes
            formation of oocysts that release into the tick’s hemocoel.   (Canis latrans) and in experimentally infected young coy-
            The oocysts are large, spherical forms consisting of a mem-  otes, it causes similar disease to that seen in dogs. Surveys
            brane that envelops multiple sporocysts in which the infec-  of free‐ranging coyotes in Oklahoma showed that 40–50%
            tive sporozoites are found.                       were infected naturally with H. americanum.
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