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Wirusy Heartland, Bourbon, Oropouche I Keystone Oraz Bornawirus Wiewiórek Różnobarwnych: Stan Obecny Oraz Perspektywy Epizootyczne I Epidemiologiczne

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Wirusy Heartland, Bourbon, Oropouche I Keystone Oraz Bornawirus Wiewiórek Różnobarwnych: Stan Obecny Oraz Perspektywy Epizootyczne I Epidemiologiczne Prace Poglądowe Wirusy Heartland, Bourbon, Oropouche i Keystone oraz bornawirus wiewiórek różnobarwnych: stan obecny oraz perspektywy epizootyczne i epidemiologiczne Zdzisław Gliński z Wydziału Medycyny Weterynaryjnej w Lublinie variabilis, a transmisja wirusa jest zarówno horyzon- Tick-borne viruses, Heartland, Bourbon, Oropouche, Keystone talna, jak wertykalna (4). Trzysegmentowy genom and variegated squirrel bornavirus: current situation and future epizootic HRTV jest zbudowany z jednopasmowego RNA o pola- and epidemiologic implications ryzacji ujemnej, przy czym segment L genomu kodu- je białko nukleokapsydu i niestrukturalne białko NSs Gliński Z., Faculty of Veterinary Medicine, University of Life Science in Lublin (39 870 Da). Segment M koduje strukturalne glikopro- teiny Gn i Gc, które wiążą się z receptorami komórki Arthropod-borne viruses have continued to emerge in recent years, posing gospodarza i są celem działania przeciwciał neutrali- a significant health threat to millions of people worldwide. Ticks and mosquitoes zujących, a segment L koduje polimerazę RNA zależ- transmit wide range of viruses to humans and animals worldwide. During the past ną (5). Białko nukleokapsydu jest immunodominantą few decades, it has been noticed that the number of reports on ecoepidemiology i u eksperymentalnie zakażonych zwierząt przeciw- of arthropod-borne diseases in humans and animals has increased. Discovery ciała skierowane przeciwko niemu nie neutralizują of new viruses Heartland, Bourbon, Oropouche, variegated squirrel bornavirus wolnego wirusa (6). HRTV wykazuje bardzo duże po- and Keystone, has raised many questions in the scientific community as it is not dobieństwo w sekwencji nukleotydów z flebowirusem clear where from these viruses come, either they are already existing viruses, or SFTSV odpowiedzialnym za zespół ciężkiej trombo- a novel species evolved from viral pathogens. The maintenance of an arbovirus cytopenii z gorączką, występujący u ludzi w Japonii in nature, involves a triad of interactions between the virus, the vertebrate host, i Chinach (7). Nie wiadomo, czy istnieje możliwość and the arthropod vector. Study on pathological complications regarding to zakażenia się HRTV na drodze człowiek → człowiek, these infections are still investigable. tak jak to ma miejsce w przypadku SFTSV, który jest obecny we krwi, w gardle, moczu i kale pacjentów (8). Keywords: arboviruses, Heartland, Bourbon, Oropouche, variegated squirrel Szczepy HRTV izolowane od ludzi i kleszczy wykazu- bornavirus, Keystone virus. ją ponad 97,6% identycznych sekwencji nukleotydów (9). Ślina kleszcza, która działa immunomodulują- co dzięki lektynom, wpływa na początkową replika- całym świecie wzrasta liczba chorych na cję wirusa w komórkach dendrytycznych skóry, wi- Na choroby wirusowe przenoszone przez sta- remię i miano przeciwciał zobojętniających wirus. wonogi. W USA w latach 2012–2016 liczba ta wzrosła Przeciwciała neutralizujące wirus występują w USA trzykrotnie. Ponadto po 2004 r. zidentyfikowano 9 no- u 64% jeleni, 55–68% szopów, 22% koni, 8% psów wych wirusów przenoszonych za pośrednictwem ko- i 4% oposów (10, 11). marów i kleszczy, a wśród nich tak groźne, jak wirus W 2017 r. w USA stwierdzono 20 przypadków za- Heartland, Bourbon i Keystone. Do nowo zagrażają- chorowań wśród ludzi. Okres wylęgania choroby trwał cych wirusów występujących w różnych regionach około 2 tyg. Chorobę cechuje gorączka 38°C, bóle gło- świata u różnych gatunków zwierząt, często o cha- wy, stawów i mięśni, osłabienie, nudności i wymioty, rakterze zoonotycznym, ale o słabo poznanych wła- biegunka, trombocytopenia (<150 000 mm3) i leuko- ściwościach chorobotwórczych – oprócz wymienio- penia (< 4500/mm3), wzrost aktywności aminotrans- nych – należą bornawirus różnobarwnych wiewiórek ferazy asparaginianowej i aminotransferazy alani- i wirus Oropouche. Te nowe wirusy stanowią najpraw- nowej we krwi, które osiągają wartość maksymalną dopodobniej identyczne zagrożenie, jakim jest od kil- w 2. tygodniu choroby. Zajęty procesem chorobowym ku lat wirus Zika (2, 3). jest ośrodkowy układ nerwowy i występuje rozsiana wewnątrznaczyniowa koagulopatia. Śmiertelność Wirus Heartland wynosi około 12% (12). W USA opisano też przypadek hemofagocytowej limfohistiocytozy (HLH) u zakażo- Wirus Heartland (HRTV) jest flebowirusem z rodziny nego pacjenta z immunosupresją (13). Wirus stwier- Phenuiviridae (poprzednio Bunyaviridae) z komplek- dzono w wątrobie, sercu, trzustce, płucach, jelitach, su serologicznego Rhanja virus, odkrytym w 2009 r. jądrach, skórze, mięśniach i mózgu, węzłach chłon- w Missouri w Regionalnym Centrum Medycznym nych 68-letniego zmarłego pacjenta. w Heartland. Występuje on w kladzie wirusów SFTS Udało się zakażenie doświadczalne HRTLV myszy, (severe fever with thrombocytopenia syndrome), łącz- królików, kurcząt, kóz, saren i chomików, ale w żad- nie z wirusami Lone Star, Sunday Canyon, Malsoor nym przypadku nie wystąpiła wiremia. Dobrym mo- i Albatross Island/Hunter Island. Wektorami wiru- delem doświadczalnym są myszy i chomiki z immu- sa są kleszcze Amblyomma americanum i Dermacentor nosupresją (14). U doświadczalnie zakażonych szopów, 616 Życie Weterynaryjne • 2019 • 94(9) Prace Poglądowe kóz, kurcząt, królików, chomików oraz myszy po- i stawów, utrata apetytu, osłabienie, nudności i wy- zbawionych receptorów dla INFα, INFβ i INFγ (AG129) mioty, biegunka, wysypka plamisto-grudkowa na wystąpiła pierwotna i wtórna odpowiedź immuno- tułowiu, szyi i klatce piersiowej. Pod koniec choro- logiczna, ale poziom przeciwciał neutralizujących by rozwinął się zespół ciężkiej niewydolności odde- HRTV był niski i tylko u myszy AG129 wystąpiła wi- chowej. Zgon nastąpił po 11 dniach trwania choroby. remia, pojawiały się objawy chorobowe i myszy pa- Charakterystycznym objawem jest limfopenia i trom- dały, przy czym śmiertelność zależała od wielkości bocytopenia oraz wzrost aktywności enzymów wą- dawki zakaźnej. Występowały wybroczyny w wątro- trobowych (21). Wirus Bourbon wykazuje duże pokre- bie, obrzęk śledziony oraz duża ilość antygenu HRTV wieństwo z wirusami Dhori i Batken występującymi w komórkach jednojądrzastych i hemopoetycznych w Afryce, Azji i Europie (22), przy czym wirus Dhori śledziony. Badania histopatologiczne wykazały tro- jest chorobotwórczy dla człowieka (23). Wirusy Dhori pizm HRTV do komórek jednojądrzastych śledziony, i Batken izoluje się od kleszczy z rodzaju Hyalomma, komórek Kupffera i komórek jednojądrzastych miąż- a przeciwciała dla wirusa Dhori występują u droma- szu nerek. Makrofagi śledziony uczestniczą w wywo- derów, kóz, koni, bydła i ludzi (24). Nadal nieznane są łaniu trombocytopenii. Choroba u małp Rhesus miała rezerwuary wirusa Bourbon i gatunki zwierząt wraż- podobny przebieg do łagodnej postaci choroby u ludzi. liwe na zakażenie oraz jego chorobotwórczość dla Występowała trombocytopenia, leukopenia, wzra- zwierząt. Uważa się, że może on stać się nowym bar- stała aktywność enzymów wątrobowych i enzymów dzo groźnym patogenem zwierząt (25). świadczących o uszkodzeniu mięśnia sercowego (15). Celem dokładnego ustalenia roli jelenia wirginij- Wirus Oropouche skiego w epidemiologii HRT zakażono 5 jeleni i mo- nitorowano objawy chorobowe, obecność wiremii Wirus Oropouche (OROV) z rodzaju Orthobunyavirus i serokonwersję. W żadnym przypadku nie wystąpi- (Bunyaviridae) wywołuje u ludzi gorączkę Oropouche, ły objawy choroby, wiremia, serokonwersja i wysiew na którą w Brazylii w ciągu 48 lat chorowało 0,5 mln wirusa użytego do zakażenia. Pomimo niewielkiej ludzi. Należy do grupy serologicznej Simbu, w której liczby zwierząt użytych w eksperymencie starano wyróżnia się 7 kompleksów serologicznych: Akabane, się wykluczyć udział jelenia wirginijskiego w rezer- Manzanilla, Oropouche, Sathuperi, Simbu, Shamonda, wuarach HRTV (16). Shuni (26). Sferyczny wirion (80–110 nm) z lipidową Do rozpoznania zakażenia wywołanego przez HRTV osłonką zawiera 3-segmentowy jednopasmowy RNA stosuje się test RT-PCR, test neutralizacji redukcji ły- (L,M,S; 27). Segment L koduje L proteinę (261.25 kDa) sinek (PRNT), testy immunosferyczne do wykrywania i polimerazę RNA-zależną, M koduje 2 struktural- IgM i IgG w chorobach arbowirusowych (MIAs; 17, 18). ne glikoproteiny Gn (28.03 kDa) i Gc (107.14 kDa) oraz białko niestrukturalne NSm (26.65 kDa), a segment S Wirus Bourbon koduje białko strukturalne nukleokapsydu (26.26 kDa) i białko niestrukturalne NSs (10.65 kDa), NSm odpo- Ten RNA wirus z rodzaju Thogotovirus (Orthomyxo- wiada za replikację wirusa w organizmie ssaków i ko- viridae) wyizolowano w 2014 r. w hrabstwie Bourbon marów, a NSs jest antagonistą interferonu (28). (Kansas, USA) od człowieka, który zmarł po ukąsze- OROV został wyizolowany w Trynidadzie i Toba- niu kleszczy (19). Wiriony o różnym kształcie – od go w 1955 r. z krwi chorych ludzi i z komarów Coqu- kulistych (średnica 100–130 nm) do nitkowatych – illettidia venezuelensis, a także z komarów z rodzaju posiadają wypustki. Genom złożony z 6 segmentów Ochlerotatus w Brazylii. Wektorem wirusa dla pta- stanowi jednopasmowy RNA o polaryzacji ujemnej ków, naczelnych, torbaczy i leniwców są komary Aedes (20). Segment 1 odpowiada za ekspresję podjednost- serratus i Culex quinquefasciatus. Serokonwersja u lu- ki polimerazy PB2, segment 2 za ekspresję podjed- dzi i zwierząt występuje w Argentynie, Boliwii, Ko- nostki polimerazy PB1, segment 3 podjednostki po- lumbii, Trynidadzie, Brazylii, Panamie, Peru i Ekwa- limerazy PA, segment 4 za ekspresję GP glikoproteiny dorze. Rezerwuarem wirusa są m. in. leniwce, małpy (~60
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