A Review of caulleryi Infection in

著者(英) Morii Tsutomu journal or The journal of protozoology research publication title volume 2 number 4 page range 128-133 year 1992-10 URL http://id.nii.ac.jp/1588/00001700/ ノ.Pr伸′二〃り/,伽.\・J./ヱ(YりJり舛甘 (1叩1項直王りウリプ.紬ピ‘J′て・ん(■川仙/ノ什〃…√馴“′=V〃/椚′J〟′/′刀′ナ川′狛叶打

AReviewofLeucocytozooncaulleryiInfectioninChi亡kens

TSUTOMU MORII

伽ブナ〟/伽抑′イP〔J仙、〟り/りg.)、,町制■玩【ノ両州吋∫‘∵加0/q/■ルわ〟f〔血ノ・ 5’ノ7~/山神乙いV血ん乱丁(勅〃ノ#/,ノ呼Ⅵ〃

Rcc由仁d20Sc匹・叩じrlリリ2/Aし■叩【d50ctol光rlリリ2

Keyword:L(,uCOCytOZOOF.CaZL]lcryl/leucocytozoonosis′, CuLjc・)ides bitingmidge

ABSll托ACT LeLLCOCyLozoo几Cau乙Leryiis an etiologjcalprotozoan of chickenleucocytozoonosis commonly fouEldin various Asian countries・Leucocy壬ozooTL C皿‖eryiis transmitted by CLLLicoides biting midges arld shows highinfeetivity and pathogenicity to chick ens.The severity of・clinicalsymf)tOmS and mortality deperld on the number of叩0 rozoitesinoculated.ScverelyinElected ehickcn50ften die of hemorrhage・∧1most allof the chiekens eha11enged with sporo∠Oites after a primaryinfection at var- ious ages show comp.ete resistanee Lo rcinfection and the aequircdimmunjtyis cxpressed against the second generation of schizogony・Solub】e antigens are found in thc sera of chickens between thelOしh and15th day after sporozoiしCinocula tion.These antigens originate from second-gerleraしion schizonts・^ntibodies against the anしigens of each developmentalstage of L.cauLLeryiare recognizedirl thc sera ofinrected chiekens.Each stageis antigenically distinct・

INTRODlfCTION ANl)HISTORY Many species of LetLCOCytOZOOn belonging to the suborder Haemosporina are kr10Wn from various kinds of throufミhouL thc world.Since LetLCOCyLo2:0071CauLLeryi as an etiological agent of leucocytazoonosis of chickens was first described by Mathis and L昌ger(1909)in Tonkinin Southeast Asia,the prevalencc of chicken leucoeytozc.onosis(==”Sed byinfection with this parasite has been recognizedir) various^sian counしri(ラS(Fallis et al,,1974).Akiba cL al.(1958)reported the f-irst case of chiekenl(1uCOCytOZOOnOSisir)」apan・ Because cIfits severeinfecLious efLfec・tS eauSing deaths due to hemorrhage and anemia,retarded growth and reduetion of egg production ratein chiekens,L.catL卜 Leryiis considered as one of Lhe Tlighly pathogenic protozoa of chickens・ ^kiba(196O)dernonstrated that L.cazLLLeryiis transmitted biologiea11y by CzLLi- coides biting midgcs be]onging to the rarnily oflL)iptera・It has beerlClarificd up to this time that most of the specics of Leucocytozoon are LビリC〔〕CY’′J一(つ∠Cブ(ラ八JCÅ描.J」~二Jn′ノINドド〔、‘1】0\1\(二川CKENS

Lran細itted by5血1血bla、ek rli執Bennett射al.(1965)p呵JOSed t山江・帥鳳雛 引率㌍iesIぬieh are transmitted by澄ま取払エ、jM b】8ek rlies are tαbeぐ1as鳥iやiedinモO t蛎色鯛uS血c8ぐツ亡児8∂氾and、tbose whieb are transmi=烏d by C址孟まcロゴd由biti噸 midge′写int8a g釧uS服伽.Al抽ou掛Suモカgenefie na血eS Were uSed ror紳me ti聡, LIsuet;al・冊73),Levine Ll射3),孔ndFaIlisotal・(197ヰ)hadequallypropo苔ed uは=鹿genusAk抽=hbuld t始a紬b脚11S When thc biologicalcharact即Of e荘Ch 報鍵ies beldnging to the f如ilyLeucoeyto署Ojdaehadyct to be jnvesti脚Led thro噸h 耶re deta‖ed印叩郎is8nS. 昆uEOeyiouon cau揖離無is noteaSily transnitt色d by bloQd pas/Sag¢極顛u如=弛壇 pr如白文Oan d(ie£nOt under酢】SehiZOg鋪yin£ireulati月gbloロd亡母=s0で頭icke舶.N蜘, e丈petiqr鐙ntalinfectic・nS With L,C皿毛Ler涙iinごhickens areinitiated by theinoeul8- ti柳川げ句紬柏ZOiteS obtained frgmi爪fecも如払エゴ、c8査dgs bitimg midges.舶riiamd Xit蜘肇(196鞄)d郡ised al鵬0柑t¢ry仁Qlonization methodぬr払‖仁0池ぎ臆噂頗卸融 珊ナyi咤a5a VeetOr f8rl.C已祉乙且訂ツ豆,馳erer8re,it beヒame p8S5ible′t0折ai机裏口 ム.亡d址去乙鐙γタ豆in thelabnrato呼and tふ鱒r晦川mbree文tend細島tudies on t血e台Ⅹp即卜 爪entalinfeeti(】nSin chickens. 蝕iekeね1肌eOCyt耶OnnQSis e射Sed byin転巴tion vithJ.,C且廿‖gγyまhag beenぷ旭ied by聯叩investigatorS,and the FeSult番てげthese studics havebeerL re/Vieや歪d由XteTr- SiYely(Akiba1970;Falliset al・、1椚4き‘This revieww=fcrcu貰1刀15tudi顔Of life呼ele,叩eCifieity,and孟mu舵re翠pO鵬esがchickens Lロム.射m右拍γ再.

【.1ドE Cl’CLE ム孤C86ダね男即花C庄址乙エe′叩盲baぷtbree酢OWth富tageS SuCh a名馬Cbiz匂gOny郡d霞am台地一 郎川yineh地色机㍉andsporo卵nyinC址‖c8畳如bi…gmid酢S・L孤C頑頗純朴頼か エ絃γツ盲inf色eti8n StartS Wh朗=細jnfeeted蝕乙壷由id鮎b′iting爪idgeしak軽写且bl瑚8 鱒須 frQm盈ehiek¢n,sj血ult8嘩Ou写1yinjeeting sp()r(〕ZOitesinto t帖bloQ鹿tre細 ロf th8油iek細(Akiba196q;胸riifさt射.119略;Moriiand Kit明細1968bト Sueわ 叩即QZ¢it昏Sinvade the¢apil18ry endp値elialeel15in the、,1un臥1iyer, and bursa of Fabricius and dcvel叩into r凱nd firSトgeEleratibn sC山王¢nt皇,血iCh m佳遣uてe and relea石倉firs仁詳meration舶一也gOiteSOn day5postinfeeti蜘(Moriヱ机d Pu如舶1爾れ舶turerir5ト脚er盆t加島ehizontSar?Ⅴ即i8bleiqsiz¢famgimgfrom 胡to6さ飢Ⅷiれdi良meter・Mature fIrst一節噂rationm即OZOite亭由由Sufi鴫7.1即in lengt且単純1叩g and slender,and aFel、eS5thanl〟min width.Tbese mer脱¢ites, i∫ltu川,iれⅤ細e the仁apillary牽ndptbelial¢ells of various tis首ueS a鵬′8r紗那8nd b餌珊elar絆SeCOnd‾等eneration露だhiz珊tS【=m曙aloschizonts)tムat r色′1eaSe点燈伽nd一 ぢ飢8PatiOn merOZ8iteちOm dayl4むr15p8Sti爪feetion(蝕iba1978;地iba蔦t al‥ 1971;Mopii白t al.,19眺).ぬture$eCOnd-generatlOn SChi紬nt写are f叫nd e離rar 軽Il叫1arlyim v8ripus o嗜anβand ti等叫eS ahd rea亡h siz守S8fりp t¢現昭ノ封鱒律d卜 如鵬t紆.Aが顔t variati6nin the si乞e of mattlre S愈COTEdgeneFatioTISehi耳QntSis 洩○叩(Eit8¢k8et al・,1g72)・Tf拇雲e紀聞d一郎那er如ion me川ZOit巳S neaSuri喝∂bQut l・7J川1JIle唱t上1andlFLmi71Vidth∴are耶aljn shape and抽vadeerythp研CyteS 湘dgr¢Winto即metα仁yte㌫抽江ure若色metQeyteS CO岬1eしely freed.from h8St eells 宅岬pe即iflthe peripberalblood of the ckick即On d8yS18-24po$tinfee曳i¢n.T短 冊a仁和g即肥tPeyteS are8匝ut15ヵ川in diameter and spheriea18r OVOidin油ape. 馳e、扇eごQg細etO野te5are丑1ittle s皿allerim size tham LhelmaCrO酔血e紬cyt色S, CuLicoiぬ兆biting midge that takやS a blood巾ealthereforeingeEtS舶ture ga舵柏一 野毛金S・The mierqgame紬qyte for耶8蓋1en鹿r雨亡r喝a腑tes(Mqri.iet al‥1墾注射. T鹿腑til¢血icr瑚am如e f即tili五官S the血erp鮮血ete tねproduee亘yg姉e.Th弓をl弛訂i組1 立y酢teS r色血盆in within the conteiltS bf′t鹿bJood mbalaild tr@EISfDrminto白Io 晰モileookiれet台S38t()60mi畑te≦丑ft即b180d meal,The ookinetesで↑耶S tJ桓biti噛 軌i、噌e rbidgtlt Walland continue developl肥nt tO for血摘und bo亡ySt$.Th台0OeyStS ne卑- き山一e at餉t即ity aboutl(I〟mifldi量腑tセr andI咋Odu亡e many SPOrOZQite5W血ieh卿卜

/-‘丁・ LEuCOCY71(〕ZO〔)N rAuJ.[J:ln/-J】NIてIiCT10NIN CHI⊂KENS grateto thc sa=varyglandsorbitingmidgein2and3daysat250C arter blood meal刷‖ba1960;Morjiet al..1965;MoriiandKitaoka19GBb;Moriiet al‥ 1984b).The岳POrOZOites MeaSuringaboutlOLLminlerlgth andl・2iLmin widthare longand slender・什ighinfectivity to chickens ofsporozoites develops during their residencein the sa】ivaryglands of bjtingmidge(Moriiat al・,19自4b)・The sporozoites enter the chicken during bj ting midRe feeding.

SPECIFICITY Le址COCytOZOOnCauLLeryiwas first observed byMathis and L占ger(19O9)in the peripheralbloodofchickens・Since this originalreport・naturalinfeetions with L.c血LLeryiinchickenshavebeen recognized byrDany【nvestigators・Mathisand L占ger(1910)kept chickens,ducks・geeSe,turkeys・andpigeonsinapenwith chickens with L.cauueryi.Onlychickensinfectedwith[′・CaLLILeryi・Moriiand XitaokE)(1971)reported that pheasants,Japarnese quails,bclb-Whites,quinea fowIs, and bamboo partridges wereinDCuLatedintraver10uSly with sporozoites of L.cauLr 乙ery.but noneof thcsebirds wasinftLCted・Theseresults suggest that thechicken is the only bost forl.c皿£乙eγy豆・ The pathogenicityof L,CauLLeryihas been studiedin experimentallyinfected ehiekens,and the results of these studies were reviewed byAkiba(1970)and Falljs et al.,(1974).Moriiazld Kitaoka(1969)reported that chickens vereiT). fectedwith a single sporozoite of L.cauueryi,arld that the severity of clinical symptoms and themortalityin chicJ{enS rOSeWiLh anincreasein the number of sporozoitesinoculatcd・Thepattern ofparasiteniairLChicJ{enS WaS dete「minedby the time and dose ofL the firsLinoculation when the birdsl柁reinoculated vith different doses of sporozoites repeatedly two or more times atlor2days,inter- vals.Cliniealsymptoms vere first observed on day12after sporozoiteinocula- tion.Chickens harbouring heavyinrections werelistless,discharged green feces, lost their appetiLe,and often died of hemorrhage・Survivors suffered from severe anen)ia due to hemorrhage and crythrocyte destruction・Petechialhemorrhages and edema were observedin chickens withinnumerable second-generation schizonts which formed thrombiin Lhe capil)aries and blood vessels of most of the organs and tissues.

TMMUNE RESPO¶SES Some of Lhe chickens prinarilyinfcctcd with sporowites of L.caLLueryiYithir1 14days of age were reinfected with sporozoites,but almost allof the chickerLS challenged with sporozoites after a primaryinfectjon at various ages showed eom- Plete resistance to reirlfection Llnd the acquiredimmunity to this parasite was expressed against seeondLgeneration schizogony(Horiiand f(itaoka1970;Moriiet al‥1986,】989). Morii(19T2)found soluble antigensin the sera of chickens oT]days10-15after sporozoiteinoculation4The highest titer of serun-SOluble antigens was recognized 2days prior toしhe peak or parasitemialThe titcr of these antigensincreased jn proportion to thc rlumber of sporozoitesinoculated and sccond-generation schizDntS formed(Morii1977).The serumSOIuble antigens originated from the schizorltS arld WCre Of ElprOteinaceous nature(Morii1974)・Precipitating antibodles agairlSt these antigens and the antigens prepared from second-gerleration schizonts began to be demorTStraLedin the sera or ehickensirlfected with L.cau乙Lerγion the17th day,and again5t arltigcns frorn secondLgeneration merozoites arLd ganetocyLes on the 2lst day after sporozoiteinoculation・The arltibodies reacted speeifica11y vith antigens prepared from L.cau乙乙eγyi.After that,the8gaJlgelprecipitaticn test has been used for the seroiJT[munOlogiealdiagrlOSis and seでCeE)izootiolo即ごalsurveys

∴川 ⊥EU⊂(〕⊂Y′’け)∠〔)(1八一 亡ノ1=J.J.ム■J汗ノ】N巨】こ(1TIし)N L\⊂=Ⅰ(’Kl三NS

Of L.cd山乙eγ再inrt、Ctionsin cれiclくenS(Fujisal(iet al.,1979;Moriiet81.,1981ト Fluoreseent antibody techniques(FuJisakiet al,,1981;Isobe and^kiba1982). COtJnterirmunoelectrophoresis(Fujisakiet al・,1980).and cnzyT”e-1inlくediJnnunO- SOrbent assay(lsobe and StlZuki1986、..1987a)for detection of antibodies to L.c皿乙乙gγヅ豆in c血iekens have been reported. Thelong-term antibody responses to L.c皿l乙eryiwere recogT)izedin chicke71S that had recovered from a prirDaryirlfeetion〔Isobe and Suzuki1987b),but a relap、5e Or recurre爪ee PherlOmenOnin these ehickens has not yet been observed. TheiTnhtlnO酢nicity of seruJn-SOluble antigens a[nd sccorld-geTLer&tion sc71izoIlt写Of L.c血まLゼγツj tロehiekens has been recognized(Morii1978;Isobe and Suzukil測鋸 Moriiet al・,1989)・EffectiveimTmnization using a killed vaccinc prepared from C血icJ(ed organs cor)taining second-generation schizonts and serumLSOluble antigens 8f L.cauLLeγyiwas achieved experih)enta】1yin chjcJ(enS(Moriiet al..199Q). ^proportion of the chickens that had beeninRunized with the vaccine survived thelethalsporozoite challenge■This protection was attributed to parti&1stlPL Pre5Sion of the development of second-generation schizontsin vaccinated chickens after sporozoite challerlge. Fupther researchis rleeded to eluci血te theimmunogeflic components of this v由仁- Cine andiEDqLun(】dominant巳Pitope、S Of each deve、lOPTDentalstage of L.cazLLLeγyi. ChickeT11etlCOCytOZOOnOSis caused byinfection with L.cauLLeryiis an economi亡al- 1yifbPOrtant diseaSe thatis presently controlled by medic&tion.Recently,r珊ains Of chem10therapeLltic agent畠inJneat and eggs of chickerlS haveled tcla reneWediJt- terestin the develop血ent Of effective v&ecines against L.cazL乙Leryi.

REFERENCES Åkiba,K、・1960,Studies on the LeLLCOCツtOZOOn foundiT)the chickens.inJapan,ⅠⅠ. C)n the transnission of L.cauLLeryiby aγakawae.JPn.].Vet.Sci.22: 309-317. 此iba,Ⅹ・197O・Leucocytozoonosis of chicJ(enS・NaLl.ITLS土.ATLれt]eaLLhQ.(]れ),10 挿画封.ノ:13ト147・ Akiba,K.,Intli,S.&1shitanl.R.1971.Morphology and distribution ofiJltPaCell11lar ぷehizontsin chickens experinentallyinfected with Akiba c血LLeryi.NaLL.I花S丈. 如血軋椚拍‖血8.(J如.ノ,11:109-121・ Akiba,Ⅹ‥ kavashima,H.†Inui,S・&Ishii.S.1958・Studies on Lg比C(】Cツ上ozロ0花Or ChickensinJapan.工.Naturalirlfection ofI..cazLL乙eryi.BtLu.NatLI7LS土.Ani 領地い用.〃如止卜仙.34:163-180(InJapanese.English sum山8ry.). Bennett,D.F.,Garnhanl,P.C.C.&Fallis/,Å.M.1965.Ur)the status of the genera土成 COCyfo2:00托Ziernann,1898and Hac舵Ot・rOtetLS f(ruse,1890(Hae7nOSP(Iridiid8:IJeuCO+ CytOヱOidae amd Rae血OprOteidae).C皿.J∴払8l.43:927932. Falliぶ,A一肌,Desser,S・S・&揖Ian,R・H・1974.On species or上胤COCツtOZβ8礼 月dり.Pqγ¢- 引圧dL12:卜67. Fujisaki,K.,Norlaila,M.&Cheah,T.S.1979.ImnunologicalsllrVey Onleucocyto▼ ZOOnOSisin chickensin Penisular .Hajian VeLeri7tar.11:3D36. Fujisaki,K.,Tak81阻tSu,什.,Kitaoka,S.,Suzuki,K.&Ⅹamio,T.1981,DirectimJ)unO- fluorescent staining of Leucocyto2:007t CatLりeryiof different deve、1opmental st喝eS.抽乙.血t.触れ加盟乙土九Q.け帥.ノ,皇1=73-79. Fujisaki,K..Takamatsu.H_,Kitaolくa.S.,Su21uki,K.&Kuniyasu,C.1988.Rapid detee- tion by counterimmunoelectrophoLleSis of antigens8nd antibodiesin the sera8f ChickellSinfected Yith LetLCOCy土ozoon ca7Lueγy査.NatL.Tns土.血irA.HeaLtfLQ.(]匝.), 28:96-108. 鮎叫C..(.,CampbeJl,G.R.&Levine,N.D,1973,^checkrlist of the species of the LetLCDCytOZOOn(,Plasnodiidae).].PγOtO2:00L.20:195L2n3.

l’、ニー J-FUC〔)rl′‖ノーり/し九)八r し■ノl=しりノく1■/l\卜=㌧、■「lいNIN(二】1JClくIiNS

Isobe,T.&^kiba,K.1982.lndirectimmunofluorescent antibody testin ehickerl lcucocytozoQnOSis.J払ぃ.J花S去.血れ鮎α乙ま九Q.り帥.プ」22:163-169. 1sobe,T.&Suzuki,K.19貼.Enzyme-1inkedinmur10SOrbent assay for detection of anしih()dy LoJg址CO印fozoo几Cα山Ieγツi.血豆α花P(ユt加且.15:199-211■ rsobe,T.&Suzukj.K.1987a.Detectit-n Of serum antibody to LeucDCytOZOOn c皿L乙eryinaturallyiTlfccted chickcns by enzyFne-1inL(edimmunos()rbeLlt aSSay・ J匝.J.レet.gc宜.49:165-167. rsobe,T.&Suzuki,E.1987b.Longterm antibody response to Le址COCyLozoo几 c血LLeryiin experimentalyinfected chickens・J匝.].Vet.Sci.49:357359・ Kitaoka,S.,Morii,T.&Fujisaki,7(.1972.Quantitative observation on schizogony Or服さbd C皿乙乙erッijn chickens・肌止L」取高.血れ〟g庄1f九Q.rJ如.プ,12:151-160・ LcviJIC凡t].1973.1)rotozoan parasites of■domestic anima】s and of man.2rld edリ Burgess,Minnesota. Mathis,C.&Ⅰ一昌ger,軋1909.J.e址CDCγf∂ZOOれdela poule.Coゆf.〟e几d.ゴビd几C.50C.β豆0乙. 67:470-472. Mathis,C.&),tLger,M・1910・Nouvelles recherches stIr Leucocytozoon sabrazesiet l,e7LCOCytO2:00n Ca7LL乙eγyidela poule doJILCStique du Tonkirl・BtL‖.Soc.PaLh.Exot. 3:504-51(1. Morii,T.1972.PreseFICe Of anLigens and antibodiesiFlthe sera()f chickens inrected with舶ibd C皿且エeγツ童.地上LJ几5と.血れJJgα乙川Q.け帥.ノ∫12:16ト167・ Morii,T.1974.Characteriz且tior10r antigens and specific antibodiesin the sera Of chickensinfected with Abiba cauLLeγyi.NatL.1nst.Anim.HeaLth(.(Jf,71.),14: 174-181. Morii,T.1978.Soluble antigcnsin the sera of chickensirlfected with L飢ICOCytO 之OO几C血乙1gγツi.PγOCggdi花gS OJ土九eJざtJ(ゆ肌gSg-Geγ爪d花C(フ〔ゆer8£室町e Sッ汀ゆ∂S盲l皿 8花Pγ0土ozoα托βise丘Sg5.211-217. Morii,T.&Fukuda,M.1992.ObscrvLltions on firsL-gCr)eratiorlSehi乙OgOny Of Leuco cジムoz叩花Cα山乙gγy童in chi亡kens・J.Pγ0£oz80エ.39:28l2きけ. Morii,T.&Kitaoka,S.1968a.T上IClaborat(〕ry eOl(】rlization of CLL己icoides arabawLZe (Diptera:Ceratopogonidae).Na土乙.Inst.AniJn.HeaLLflQ.(Jt)7t,),8:204-2()9. Morii.T,&Kitaoka,S.19朋♭.Tnfluence〔)f temperature on the sporogony of catLLLeγyiin three CuLicoides species・NalL.Inst.Anim.tleaLthQ.(Jt・n.),8:210- 21〔;. Morii,T- &KiLaoka,S.1969.Susceptibility of chickens of different ages to sporor ZOites or劇沌αC瑚1乙gγツ童at Various doses・帖t上.J花ぶと.血れJJeα乙f九Q.〔J♪札J, 9:104111. Morii.T・&Kitaoka.S・1970・Some aspects oninmunity Lo^biba cazLueγyiinfection in chickens.肋亡乙.上靴S土.血れ〟e瓜と壬九Q.(J匝.ノ,10:15卜Ⅰ59. Morii,T.&Kitaoka,S.1971.Susceptibi=ty of the gal】inaceotlS birds to c皿1乙eγyゴ.他壬い几雨.加血軋肌氾‖h Q.〔J帥.ノ,11:17什」7l. Morji,T・,1ijima,T・&f;ukuda,M・1984a・Fine strucLure of Leucocytozoon ca7Lueγyi during microgamt〕tOgeneSis and fertili7atior]・Zbl.Bakt.Hyg.^.256:314-322. Morii,T・,Kitaoka,S・&Akiba,K・1965・S(】me 王nvesLigations on tlle SPOrOgOny Or Leucocyto2WOn Cau乙Leryiin7ab【1ratOryrreared biting midges of four CuLicoides SpeCie5.肌止L」厄封.血れ〟e正己と九日.りp几.ノ,5:109-110. Morii,T・,Matsui,T・,Ii]ima,T・&Fu]inaga,F▲1984b・tnfectivity of LezLCOCyfozocm cd址工乙grッ盲 SpOrO別再tes deve】opedi花封‖ro and豆托封宣む0.ノ花土.ノ.P皿γ瓜5卓上81.14: 135-139. Morii,T.,Mat$ui,T,,Xobayashi,F・&Iijima,T.1g89.Sone aspects of Le比COCylozoon c皿乙1eγヅ盲 reinreetionin chickens.P(王γ丘Si土Dl.斤g5.75:194-198. Morij,T.,Nakamura,K.,Lee,Y.C。,Iijima,T.&HoJi.X.1986.ObservaLiorlS On the Taivanese sL「ain orJe址C()Cツl8ZPO几C血乙エeγγ豆(Haemo5POriIla)in ebickens.

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J.Pγ0土oz8Pエ.33:231-234. Morii,T.,Fujita,].,Akiba,X.,Isobe,T,,Nakamoto,Ⅹ..Masubt]Chi,K.&1shiharalH. 1990.Protectiveirnblunity to Leucocy10ZO血CauLLeγyiin chickens by a ki11ed v鱒Ceine.PαγαSjtol.尺e5.78:6ユロー632. Morii,T.,Shiihara,T,,Lee,Y.C.,Mar)uel川.F.,Nakamura,.K.,上ijih)a,T.&HoJi.K. 1g81.SeroimnTunOlogicaland parasito10gicalsurveys of Leucocyto2:00n C此乙ZeγJ7i infeetionirlChickensin severalÅsian cour)tries.Jn土.].Parasi土0乙.11:1B7-19D.

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