PSG. TS. PHAM VAN TY RVI en HOCPGS. TS. PHAM VAN TY VIRUT HOC (Sach ding cho sinh vién ngénh Khoa hoc tu nhién) NHA XUAT BAN GIAO DUCI Oi gibi thiéu Virut li mét loai vi sinh vat rat dac biét. Ching rét. nhé bé, obi cé thé quan sat due duéi kinh hién vi dién ti Ching lai chua cé cia tric té bio nén bit buéc phai ky sinh ndi bao trén co thé’ mét sinh vat khac ; c6 thé’ké' déh virat cda vi khudn (bacteriophage), cua xa khudn (actinophage), cda vi khudn lam (cvanophage), cia ndin (mycophage), cia tao (phycophage), cua thuc vat (plant viruses), cia dong vat nguyén sinh (protozoal viruses), ctia déng vat khong xwong séng (invertebrate virases), cia cén tring — (insect viruses) vd cia dong vat cé xuong song (bao gém cd nyudi) (vertebrate viruses). Ching cht cdi tao bdi mét vé protein bao boc bén ngodi mét loi axit nucleic. Ching 06 thé’ két tinh thank tinh thé’va ton tai ldu dai nur mét hod chat. Tuy nhién, ching 6 ddy du cac hoat dong séng nhu di truyén, bith dj, tang trading; sinh sin, trayén nhiém.., mac déu cdc hoat dong nay déu phdi gan lién vdi t&’ bao vat chu. $0 lodi virut I rat 1dn va chia 65 thé thong ké Aét duoc. Ching han riéng cdc virut thuc wat hién dé biét én 1000 Jodi, Nam 1960, Uy ban quéc t& phan loai virat International Committee for the Taxonomy of Viruses (ICTY) dupe thanh lap, tir dé dén nay di cé nhiéu cuge thdo Iudn va di cé duge mot he théng phan loai khd én dinh. Nam 1990, ICTV di xde dinh dug 20 nguyen téc ding dé’ dinh tén virut va nam 1995 da dua ra hé thong phan loai cdc virut déng vat gdm 23 ho hdc nhau, Nim 2000, ICTV dua ra hé thdng phan loai virut thuc vat bao gdm 15 ho va 73 chi.. Nhan logi quan tam rat nhidu dén virut trade hét Id vi tinh nang gay cic bénh truyén nhiém cho nguoi, gia sic, gia cam, cay wong, cdy ring va sit dung trong céng nghé lén men... Ching ta déu biét dé méi de doa ctia v6 sO cic bénh hig nghéo dle virut gay ra 0 ngwoi (bénh dat, viém nao, viém ming nav, viém gan, quat bi, tiéu chay, sot vang, cdm Janh, cim, diiu maa, thuy dia, zona, herpes, viém phdi, sot xudt huyél, ung thi bach cdu, SARS, AIDS va gdn day nhat hi nguy co lay virut HSN] tr gia cém sang nguéi...). Nganh nong nghiép cing hét stic hing ting trade cic bénh nghiém trong do virut gay nén ma chia cd thadc phong tr dic hiéu (bénh khdm 6 thude li, du dé, khoai fay, dura chudt, ..., bénh xodn Ii ca chua, bénh vang Ini dIéa,...). Cong nghé lén men nho vi Khudn dé tao ra thud khang sinh, sin xudt bt ngot lizin, enzym, dung moi hitu co, axit hire co, thudt tri s4u BT... cung Iudn judn bi de doa boi thuc khudn the (bacteriophage). vi sinhTrong nhiéu ném qua, ngénh Virut hoc (Virology) & mide ta da cé nhéing bie tith bo ding ke. Ching ta da dti kha nang: phn lap vé xdc dinh cde virut gay bénh, da sin xudt duge rat nhiéu loai vacxin chong virut, ké cd cdc vacxin ti 08 hop nh vacxin phong chéng viém gan B, vacxin cuim,... Vic nghién citu chan dodn va phéng trénh cdc bénh virut cho gia stic, gia cdm, cho tom, c4... cing a3 cé dude ning thanh tau khong nbd. Tuy nhién, cic tai liga gido khoa vé Virut hoc & nuiie ta con rit hiéin hoi va vi vay kim han chéhiéu biét vé virut cda cdc cin bo khoa hoc, cin b6 ky thudt lim cng tic cé lien quan dén virut (phéng chéng bénh trayén nhiém, thi y, bdo vé thuc vat, céng nghé Ién men nhé vi khudn va xa khudn...). Cu6h sich Virut hoe ctia tic gid Pham Van Ty dip ing duoc nhu du néy cia déng déo sinh vién Y khoa, Sink hoc, Néng nghiép, Thus! sdn va cic cin b0 lim viée trong cic nganh ndi trén, Sich due viet trén co 30 cde tai liu cép nat va duoc trink bay mét céch dé hiéu vi nhiing tank anh minh hoa rét sing sta. Voi hy vong dua dupe kigh Unie vé virut hoc déh 86'dong sinh wién vd ban doc rong Tai, (Gi vui ming dug git thiéu cudh sich dude viét rat nghiém bic vi rat bO ich nay. Ha Noi, maa Thu ndm 2004 wy GS.TSKH. Hoang Thuy Nguyén Chi tich H6i Vi sinh vat hoc Viet Nam[A noi _déu Virut li ky sink n6i bao bit bude, cé khd nang gay bénh d moi co thé’ song tr vi khuan dén con nguéi. Trong lich st lodi ngudi, sO nguoi chét trong nhiing trn dich do virut gay ra con Jin hon tét cd cde cudc chiéh tranh, xung d6t sac thc, ton gido, nan déi, cdc vu dong dat, Id lut va tai nan giao théng cong Iai, Nhan loat ludn bang hoang trubc c&e tran dich do virut gay ra. Nim 1918, dich cim da lim cho phan mia dan s6 dja cla mac cup di 20 tri¢a ngwbi. Thuc té'cho thay ti nhing thap kj cub thé ky XX tré di ngay cang xudt hién nhiing bénh virut méi la 3 ngusi va gia stic, de doa cade séhg loai ngudi. Dai dich HIV va hang loat cdc bénh mdi khac nhir SARS, ctim gi HSNL Ebola, Marburg, Hanta,... ludn la ndj dm anh d6i véi loa nguéi. Virut gay benh 3 dong vat da khidh ngwoi ta phai giét bd cd din gia sic, gia cém, gay thiét hai lin cho nganh chan nudi. Virat gay bénh 3 tom fi nguyén nhan khién cho nhiéu ao tom chét hang loat. Virut gay bénh thue vat cling la Uni pham gay that bat mia mang. Véi nhting If do trén, viéc nghién citu virut luén duoc dy manh va da tro thanh mot trong nhting nganh khoa hoc quan trong hing déu trong Sinh hoc, Y hoc, Nong nghitp va Nu6i trong thuy sén. Cé thé néi nifa sau cua the ky XX, virut hoc dé cb nhiing bude tién khéng nging va da thu dupe nhing think qud Ién lao. Nhiéu virut gay bénh moi di dure phan lip va tim hiéu ban chat sinh hoc. Nhiéu loai vacxin thehé mdi da trén cdc kF thadt cia cong nghé sinh hoc dé ra déi, nhiéu bién phap tri liu méi cing duoc dp dung. Vide xod bd vinh vién bénh dau maa vie nd 1977 va sdp ti la bénh bai liét quy md todn cdu, dude cof li thanh tru vidai ctia Y hoc. Mat khde, do cé cit t20 don gidn va do cé hé gen nhiéa kia v6i co ché tdi bin khéc han céc co ché'don béo va da bdo nén virut duge chon lim m6 hinh Iy tudng dé’ nghién ctiu nhiéu co ché sinh hoc & mtic phan tu, din dén cudc cach mang sinh hoc cin dai véo Glita thé’ ky XX - Sinh hoc phan tt, Di trayén hoc phan tu. Do dé, n6i dung gidng day mén virut hoc dai cuong vé thuc chat ciing li gidng day vé sinh hoc phan ti - mot doi hoi cia thoi dei. Mon virut hoc vi thé cing can duge nang cao hon. O Viet Nam, Virut hoe chia thye su phat tridn dé trd thanh mot nganh nghién cite déec dp. Tay nhién nhiing nghién ciu te té vé cic virat gay bénh cling dang duoc tién 3hanh tai cdc Vién Vé sink Dich t& hoc Trung wong va Thanh pho Hé Chi Minh, Vién Cong nghé Sink hoc va mét so trubng Dai hoc Nong nghiép. Cudh sich nay nhdm giéi thiéu cho sinh vien, nghién cia sinh, vi caic cin BB Khoa hoc 6 lién quan nhiing kith thiic co bin méi nhdt vé virut hoc trong vai chuc nam gan day. Mac dit khéng phii ld cubh sich chayén khoa y hoc, nhung vi mong mudh cubh séch hit ich hon dot voi déng dio ban doc, nén doi vei cdc virut cé tm quan trong déu c6 trinh bay vé dic didm lam sing, sinh bénh hoc, dich hoc va cdch phong chong. Toc d6 phit trich ctia Virut hoc rét nhanh chéng, trong khi tém nhin cda téc gid lat con nhiéu han ché, do vay nhiing khiGin khuyét trong cudh sich nay Ia khOng thé trénh Abdi. Tic gid xin chan thdnh cdm on vé mong nhin dupe su gép 9 cia d0c gid xa gan. Hé N6i, mia Thu ndm 2004 TACGIAcAc cut vik TAT AAV Adeno associated virus (virut két hop adeno) AIDS Arquired immune deficiency syndrom (hai ching thiéu huc mién dich mac phai) ADCC Antibody dependent celi mediated cytotoxicity (gay doc té bao qua trung gian té bho phu thuge KT) CD4_ Clusted of differentiation (thy thé CD4) CF — Complement fixation (cé dinh bé tha) CMV Cytomegalovirus (Virut cytomegats) CPE Cell pathogenic effect (higu ing huy hoai té bao) c-one Cell oncogene (gen ung thie cila t€ bio) DTB ai thuc bao DUMD Dép ung mign dich DVCXS Déng vat c6 xuong sng DVKXS Dong vat khong xuong sng EIA Enzyme immunoassay (thi nghiém MD gan enzym) ELISA Enzyme linked immunsorbent assay (ky thuat halp phy mién dich gn enzym) Env Envelope (vo ngoai cita virut) EF — Elongation factor (yéu t6 kéo dai) EBV Epstein-Barr virus (virut Epstein-Barr) IF Immunofluoresence (mign dich huynh quang) gp — Glycoprotein HAV, (HBV) Hepatitis A (B) virus (virut viem gan A, B) HCV Lepatitis C virus (virut viem gan ©) HCoV Human corona virus (virut corona & nguéi) HDV, HEV, HGV Virut viém gan D.E, G Hl test Hemaglutination inhibition test (thir nghiem tte ché ngung ket héng céu} HIV Human immunodeficiency virus (virut HIV) HLL test Hemolysis inhibition test (thit aghigm tfc ché tan méu) HPV Human papiloma virus (virut papiloma & ngudi) HSV Herpes simplex virus (virut Herpes simplex) HTLV Human T lymphotrophic virus (virut gay benh te bao tT 6 ngudi) IFA Inderect fluorescent antibody assay (ht nghiém khang thé huynh quang gidn tiép) IFN Interferon IL Interleukin KN Khing nguyen KT Khing thé LA Latex agglutination (ngung ket hat Latex) LCMV Lymphocytic chonomeningitis virus (viem mang nao mang dém lympho bo) LTR Long terminal repeat (doan lap lai dai dau chuéi} MD Mién dichMHC Major histocompatibility complex (phite hyp hoa hgp mo chinh) NST Nhiém sac thé PHA Passive hemaglutination (ngung két héng cu bj dong) PrP Prion protein (protein prion) PrP, PrP® Protein prion cua té bio (c=celular), protein prion gay bénh (sc=scrapie) RF Replicative form (dang sao chép) RI Replicative intermediate form (dang trung gian sao chép) RSV_ Rous sarcoma virus (virut sarcoma Rous) RT-PCR Reverse transcription PCR (PCR - phién ma ngugc) SARS Severve acute respiratory syndrome (hoi chimg viem dutmg ho hap caip) Te Cytotoxic T cell (68 bao T doc) TCR T cell receptor (thy thé t€ bao T) Ty Helper T cell (té bao T tro gitip) TK Timidin kinaza TMV Tobaco mosaic virus (virut kham thuée 14) TNF Tumor necrosis factor (yéu 16 hoai tir ung thu) WHO World Health Organization (16 chitc ¥ té Thé gidi) TTK Thé thuc khudn Uy Long sequence unique (doan dai duy nhaty Us — Short sequence unique (doan ngain duy nhat) RdRp RNA dependent RNA polymerase (ARN polymeraza phy thuge ARN) VZY Varicella-Zoster virus (virut Varicella-Zoster (mot loai virut herpes)A - PHAN DAI CUGNG Chuong 1 NHUNG KHAI NI&M CO BAN VE VIRUT 1.1. VAI NET LICH SU Mt thé ky trude day, virut duge coi 1A mét chat déc mo hé lu6n gan lién voi benh tat. Thuat ngit "virut” ciing bat nguén tw tiéng Latinh "Virus" ¢é nghia 1A chat doc. Vao nam 1884, Charles Chamberland - mot cong téc vién cla Vién Pasteur Paris da ding mang loc bing sit dé téch cdc vi khudin nhé nhat va vao nam 1892 nha nghién ctu bénh hoc thuc vat ngudi Nga Dimitri Ivanopski da si: dung mang loc Chamberland dé nghién cttu bénh kham thuéc 14, Ong nhan thay dich ép 14 cay bi bénh da cho qua mang Igc van 6 kha nang nhiém bénh cho cay lanh. Mac dit khong nhin thay bat ky vi sinh vat nao nhung 6ng cilng thong béo rang chinh “virut qua loc" [a tac nhan gay bénh. Thoi bay gid nhan dinh nay c6 § nghia quan trong, bdi vi cdc nha khoa hge van chia biét duoc ‘mot t4c nhan gay bénh ndo nhé han vi khudn. Nam 1898, Martinus Beijerink lap lai thi nghigm cla Ivanopski va nhan thay virut 66 thé qua duge ban gel va hoat lye van con trong id va dat kho nhung higu lye bi mat nhanh chéng khi dun sdi. Ong két luan day 1a "chat doc séng” (contagium vivum fluidum) ché khong phai 1a vat thé ran. Cong vao nam ay Paul Frosch va Friederich Loffler thong bao bénh 1 mém long méng 6 ngua ciing do virut qua loc gay nén. Nam 1901, Walter Reed vi nhém céng tac cia 6ng & Cuba phat hién téc nhan gay bénh sét vang cing 1a virut qua loc. Sau 46 cdc nha khoa hoc dA tim ra mOt sé loai virut, vi du virut g4y bénh dai, virut dau mia khong dé gi chui qua mang loc do vay cdc téc nhan gay bénh chi don gidn goi I “virut", Nam 1915, nha vi khudn hoc ngudi Anh Friderich Twort va nam 1917 nha khoa hoc Phap Felix d'Herrelle thude vién Pasteur (Paris) da phat hién ra mot dang virut dac biet ky sinh trong té bao vi khudn phd huy vi khudn va dat tén 1a "Bacteriophage", cé nghia 1a thé thyc Khudn hay goi tit 1a phage. Thdi ay D’Herrelle tin ring c6 thé ding thé thyc khudn dé di¢t vi khudn nhiém vao co thé nhung ahiing khdm phd sau dé cho thay thé thyc khudn c6 tinh dac higu rat cao déi voi vi Khudn ma ching ky sinh va chting bi loai khoi co thé rat nhanh chéng. Do tinh dic higu nay, ngay nay ching duge sit dung nbu 1 cong cy quan trong trong nghién cifu bién nap va nhan dang vi khudn. Nhiéu nam tréi qua, danh séch cde bénh do virut ngay cang dai. Nhitng nghién cite virut da tr nén dé dang hon khi ngudi ta phat hign ra cdc bién phép nudi chting trong ph6i ga. Vi nhan lén duoc trong phdi ga nén mac di cé kich thuéc v6 cing nhé bé virut 9van duge qui quyéi cho ring ki phiin tir song. Nhung vao nam 1935 Wendel M. Stanley thuge Vien Rockefeller 4 dua ra mot thong béo gay chin dong du luan ring Ong da két tinh duge virut khdm thude 1d. Digu nly (ang them méi hoai ughi vé ban chat song cia Virut. Phai chang mét sinh vat séng cé thé 1a tinh thé ? Vao nhitng nam 40, céc nha nghién cru da dua virut hoc dén ky nguyén mdi do hai tién bo ky thuat trong thoi ky nay, D6 la kinh hign vi dign tir va nudi cay virut trong binh nudi cay mo té bao. . Nhu vay, nho mot thiet bi k¥ thug mang Igc da dem lai khdi niém ban dau vé virut va sau dé nbd mot thiei bi k¥ thuat khac - Kinh hién vi dien tit (v6i chim tia dién tr cd do dai bude séng cue ngan, fam ting do phan gidi va tang kha nding phéng dai) di quan sat dugc virut, tim hiéu duge ban chat va chite ning cita ching. Tit vic nudi thanh cong virut trong binh da ché duge vacxin chéng mét s6 bénh virut. Nam 1952, A. D. Herrskey 4 chimg minh vat chat di truyén 6 thé thuc khudn Ta ADN. Nam 1955, H. L. Fraenkel Conrat lip rép thanh cong léi axit nucleic vao vd capsit cita virut khim thuéc Ié vi 10 ném sau, nam 1965, Spiegelman lap rép thanh cong chudi ADN cita thé thyc khudn @X174. Nam 1970, Q. Baltimore vi H.M.Temin phat hién enzym phién ma nguoc 6 virut Retro chita ARN chuéi don, Cac nghién cfu vé virut ngay cling thu duge nhiéu thin tuu méi mé gép phdn quan trong vao viéc day ii nhiéu bénh virat & nguvi, déng vat nudi va cay tréng déng thoi g6p phan vao su phat trién chung cia sinh hoc hién dai. 1.2, HINH THAI VA CAU TRUC CUA VIRUT Virut la te nha nhé nhdt 06 thé gay bénh & vat thé sng. Kich thude cita ching gidi han tir 250nm (460i v6i virut pox) dén 20nm (d6i voi virut parvo). Nhu vay virut 1én nhat cO kich thuéc xap xi kich thude cia té bao nhé nhdt mhu Chlanydiae va Mycoplasma, con virut nhé nhat lai cé cing dudng kinh tong ting v6i phan we ADN. Néu so vi E. coli thi virut 1én nhit ciing chi c6 kich thuée bang 1/10 con virut nho nhat chi 6 kich thuéc bing 1/100. Tal ca cde virut déu 66 edu tg0 rat don gidn, bao gém hai thanh phan co ban 1a axit nucleic & ben trong goi 1a J6é (geztont) va vO protein bao boc bén ngoai dé bio ve genom goi la capsit. Phac hop gém axit nucleic va capsit duoc goi IA nucleocapsit. Mot s6 cbn €6 vd bao boc xung quanh vo capsit goi la »d ngodi (envelope). Virut hoan chinh con duoc goi 13 virion va tit day chung ta dig hai thuat ngit virut va virion nhu nhau. 1.2.1. Genom cua virut Virut chi chita ADN hodc ARN ma khong bao gid chita ca hai logi. Digu nly ngugc lai v6i céc co thé c6 célu tao t& bao — trong méi t€ bao bao gi’ cing c6 ca hai loai axit nucleic. Hon nifa genom 6 co thé bac cao, cd J dong vat vA thyc vat bao gid cing la ADN chudi kép. Cdn genom, ciia virut c6 thé ia ADN hoc ARN 6 dang chudi don hoac 10chudi kép. Ca 4 loai genom ADN chuéi kép, ADN chudi don, ARN chudi kép, ARN chudi don déu tim thay trong sé cdc virut dong vat, thuc vat va vi sinh vat. Trong té bao bi nhiém, genom virut duo phién ma, dich ma dé tao thinh céc protein cau tric (protein tao capsit), protein khong cau tric (cdc enzym can cho sao chép) va tao cdc bin sao genom (axit nucleic 16i) ding cho lap rép céc virut mdi. Kich thuée genom rat khdéc nhau tir 3500 nucleotit (nhu d phage nhd) dén 280000bp (base pair), vi du & virut herpes 14 560000 nucleotit. Trinh ty cla genom virut phai duge gidi ma nbd té bao vat cha nén céc tin higu diéu khién nim tren genom phai dugc cic yeu (6 cita t€ bao nh4n biét. Genom virut rat nhé, do dé trong qué trinh tin hod dé sir dung t6i da tiém nang gen cua minh, 6 virut 6 hién tugng cdc gen chéng [én nhau va cat noi mARN [a rat phé bién. Dac diém cia genom virut Ia: + Genom ADN kép thuéng 6 virut c6 kich thuée 1én nhat, genom ADN mach don thudng ¢ virut cé kich thuée nhé hon. + Genom ARN kép tat ca déu phan doan. + Genom ARN don duge quy dinh 1a genom (+) va genom (-) dya theo tinh nucleotit cla MARN. + Phan Ion genom ARN mach don Ia genom khong phan doan (rit virut cam va HIV). + Genom cita Hepadnaviridae duoc téng hop trong té bao gan thong qua ARN trung gian. 1.2.1.1. Genom ADN ctia virut ADN cia virut c6 thé 6 dang vong, xoan (virut papova) hoac dang thang, chudi kép (vitut herpes, adeno). ADN phién ma trong nhan va duoc kiém soat béi h¢ enzym cila bao. Gen ma hod cho protein nim & cde vi tri khée nhau trén ADN, © virut adeno cde gen nay nam trén cé hai mach. GO hai déu mach thuéng 6 trat tu thang lap lai va trai chiéu c6 thé bat cap véi nhau Iam cho phan ti cé dang vong. Ham lugng guanosin va cytozin trong ADN dao dng ti 36% & virut pox dén 70% & virut herpes. Nhiéu ADN va mot s0 ARN cia virut cdn chia céc trinh tu nucleotit "thita” (khong Jam nhiém vu m hod) 6 dau chuéi. Sau day 1A mot sé vf du vé genom ADN clita céc virut. 1.2.1.1.1, Virut e6 ADN kich thude lon Genom cia cae virut thudc ho Herpesviridae 06 kich thude thay déi tir virut herpes simplex va varicella zoster (120 — 180kbp) dén virut cyromegalo va HHV - 6 (180 ~ 230kbp). ADN ma cho hon 40 protein cia virion va hon 40 protein khong cau tric (chi thay & trong té bao nhiém). Cau tric ADN thay déi it gitta c4c thanh vién cila nhém. Ching 18 loai duy nhat chita cdc doan déng phan (isomer) ciia cing mot phan tit. Genom cia herpes simplex chita hai phan n6i voi nhau bang lién két cong hod tri, mot 1ving dai duy nhat (U,) va ving ngén duy nhat (Us), d hai ddu méi ving 1a céc doan lap lai trai chiéu. Céc doan lip lai nay cho phép sip lai cau tic olla cée vig duy nhat va do vay tao ra 4 déng phan cé chitc nang giéng nhau (hinh 1.1). Genom virut herpes cing chita nhiéu doan Lap lai (muitiple repeated sequences). Cée doan nay rat khdc nhau & cée virut khdc nhau lam cho genom thay déi rét nhiéu (én hon hoac nhé hon so véi kich thuéc trung binh). Genom ctia virut adeno nhd hon, khoang 30 — 38kbp. Mi virut chita 30 — 40 gen. Trinh ty tan cing cia méi soi 1a mét doan lap trdi chigu khoang 100 — 180bp va do dé cée chudi don bién tinh 6 thé két hgp tao nén cau tric "pan - handle", dé 1 céc chudi dai va manh néi v6i nhau thanh ving va déng vai tro quan trong trong sao chép. G dau ‘' cia genom cé gin mot protein 55kDa hoat dong nhu 14 doan méi dé téng hop soi ADN mdi (hinh 1.2). Diéu dang luu ¥ 1A trong khi & viru herpes, méi gen cé mét promoter riéng thi & virut adeno genom cé cic cum gen, méi cum cé mot doan khdi (promoter). au chung Hinh 1.4, MOt sé genom cia virut Herpes. (Herpes simplex) bao gém hai doan néi véi nhau bng lin két céng héa tri, doan dai duy nbat U, va doan ngn duy nhét Us. Shai déu mi doan lai duge ndi voi cac doan lap lai tai chiéu. Caich 18 chit nay cho phép tao ra 4 dang genom khac nhau. (Theo Cann, A, Principies of Molecular Virology, 1993) Protein Gen sém Gen mugn ae amen tan ina raven strain Protein u a genstm —gensém gen sém ‘ancing 0 5 10 15 20 25 30 3538.30 kbp Hinh 1.2. Céu tao genom virut adeno. (Theo Cann, A., Principfes of Molecular Virology, 1993) 1.2.1.1.2. Cae virut cé ADN hich thitde nho Genom virut parvo 1a ADN mach don, dang thang, khong phan doan kich thudée khoang 5kb. Da s6 cc soi ADN trong virion 1A soi (-). Genom chi gdm 2 gen, gen rep ma cho enzym ding cho phién ma va gen cap ma cho protein v6. Cudi cla genom c& cdc doan xuéi ngugc nhu nhau (palindromic sequences) dai khoang 115 nucleotit tao thanh cau trdc dang cap t6c (hairpin) ding cho khdi déu sao chép. Virut polyoma 6 genom la ADN mach kép (hinh 1.3), khép vong chtfa khoang Skb. ADN 6 dang siéu xoan va lién két vdi 4 histon ca 1é bao (H2B va H4). Virut c6 6 gen do sit dung cd 2 mach ADN dé mi hod va sir dung gen chéng lép (gen ng géi lén gen kia). 12Hinh 1.3. Cu tao phic tap ciia genom virut polyoma a do thu gon cdc théng tin di truyén vao cing mét doan tuong déi ngén (Theo Cann, A., Principles of Molecular Virology, 1993) 1.2.1.2. Genom ARN ciia virut 1.2.1.2.1. Genom ARN trong virut ARN khong phan doan Virut ARN thudng cé genom nhé hon genom ciia phin 1én virut ADN, ching dao dong tir 30kb (coronavirus) dén 3,5kb (phage QB). Ching c6 thé 1a sgi (+) hoac (-). Da s6 (khong phai tat ca) genom ARN (+) déu cé mi dé bao vé dau 5' cia ARN khoi su tén cong cla photphataza va céc nucleaza khdc déng thdi xtic tién chic nang mARN khi dich ma. Doi v6i phan 16n virut picorna, mii duoc thay thé bang mot protein nhé goi 1a VPg. Dau 3' cla da sé genom virut chudi (+) cing duge gan dudi polyadenin ~ polyA giéng nh mARN 6 eukaryote. Virut ARN (-) thudng cé genom I6n hon genom ARN (+). Cau tao genom cha mét sé virut ARN duge thé hién 6 hinh 1.4. Picoma SUTR Protein Protein khéng UTR , cl do cd tic 3 5 lea oy.) Toga SUTR Protein khong Protein gure s cfu tric cau tric @ Se 09 Fla “suTR Protein Proteinkhong gute : cautic edu ruc 43 Corona Protein khéng, a Su tric ee nl AAI Protein cu tric Pramyxo 3 NS(M1) NS(M2) Hinh 1.4, So 46 genom ciia virut ARN cho thay su phan bé cla céc gen ma hoa cho protein cau tric, protein khéng cu triie, cng nhu cac ving khéng dich ma UTR (unstranslated region). 13ARN cla virut c6 thé & dang mach don hoac mach kép, thang hose khép vong. Vi du virut thyc vat luén chifa ARN chudi kép dang vong trong khi ARN cita virut dong vat di [a chudi kép hay chudi don déu & dang thang. Mot s6 virut chita genom 1a ARN phan doan, tis hai doan & virut arena dén 11 doan 6 virut rora. Viec ARN mhiéu doan va mach kép di Lam tang kh ning ma hod ctia genom vi kich thuée cia ARN bi gidi han béi co che gin vio riboxom. Sw phan cuc, Tay thugc vao chic nang cia ARN mach don ma chia genom ARN cua virut thanh hai nhém. ~ Nhém 1 g6m ARN cé trink tie nucleotit gidng véi trinh ue cia MARN, c6 du gan "mii" dé riboxom nhan dién mot céch dic higu nén cé kha nang hoat dong nhu [a mARN tic la thong tin di truyén trén genom duge sit dung truc tiép dé téng hop protein ma khong ciin giai doan phién ma. ARN nay duoc quy uéc la soi duong (genom duong). 6 cific virut 6 genom (+) khong ¢6 su khdc nhau gitta phién ma vi sao chép genom. ~ Nhém 2 gdm ARN c6 trink tu nucleotit ngugc (bb sung) vi trinh te mARN nén khong thé hoat dong nhu mARN. ARN loai nay dugc quy uée 14 soi am (genom 4m) (hinh 1.5). Ching duoc ding lam khuon dé téng hop mARN. Mu6n tign hinh sao ma wude hét phai tng hop sgi khudn ARN(+) trung gian, ri tiv soi nay tong hop nén sgi (-) 1 genom cita virut. R6 rang & day sao chép genom va phién ma 1a hai qué trinh khéc nhau, ké cd chigu dai phan tir ARN duge to thanh clng khéc nhau. Khi sao chép sé duge sgi ARN c6 chiéu dai day dé (full length RNA) tuong duong vdi chiéu dai cua soi me, cdn khi phién ma sé rao ra cdc phan th ARN ngdn dé rdi tu dé dich ma thanh cdc phan tir protein véi chttc nang khac nhau. Virut reiro la trudng hop dac bist, ching chita ARN mach don, (+) va sit dung enzym phién ma nguoc cia minh (ADN polymeraza phu thuge ARN) dé chuyén thong tin tir ARN sang cADN sau dé tao ADN soi kép dé pan xen vo genom ciia t& bao chit dudi dang provirut va tn tai lau dai gidng nhu thé thuc khudn tiém tan. mARN 5..GAC UCG AGC...3° ADN(+) 3..GAC TCG AGC...3° ADN(-) 3..CPG =AGCTCG...3' ARN(+) S..GAC UCG AGC...3" ARN(-) 5...CUG AGC UCG...3' Hinh 1.5, So sanb tral ty axit nucleic. Mach axit nucleic trong genom dude goi la (+) néu 6 {rat ty nucleotit giéng trat tyr clia MARN, con goi la (-) néu e6 tral ty nguge lai vai MARN. 1,2.1.2.2, Genom ARN phan doan Binh thudng genom ARN 1a mot sgi lién mach nhung trong mot s6 trudng hop lai bao gém nhigu doan ngén goi la genom phan doan. Vi du, virut ciim cé 8 doan ARN don (hinh 1.6), virut reo chita 10 — 12 doun ARN kép. 14Genom phan doan 1a genom chifa hai hay nhiéu phan tit axit nucleic khéc nhau vé tinh chat vat ly, ciing dugc géi lai trong mét virion. Uu diém cua genom phan doan 1a thong tin di truyén chtéa trong cdc sgi c6 kich thuée nhé hon nén it bj phd vo din dén lam sai léch thong tin. Sy sai Iéch nay rat nguy hiém d6i v6i genom ARN In. Tat nhién, virut phai c6 mot co ché boc g6i rat ky luGng sao cho tat cd cdc phan tir déu duge bao géi trong virion. Hién tuong genom phan doan lam ngudi ta lién tuéng dén NST 6 co thé bac cao. Néu mot phan tir bi hu héng thi cing khong anh huéng qué nhiéu dén ca qué trinh nhan lén cia virut. Virut ctim 1a mot vi du (xem phan virut cim). PB2 PBI PA Gen 4 [ 1 Gen2 [ 1 Gen3 1 HA NP NA Cond fe) Gen S) Gee eed Gengdi 3) MaMa NS1/NS2 iGeni7( Gana) Gens Co Hinh 1.6. Genom ARN phan doan é virut orthomyxo 1.2.2.V6 capsit Capsit 1 v6 protein duge cau tao béi cae don vi hinh thai goi 1a capsome. Capsome lai duge cau tao tir cdc don vi cu tric goi la protome. Protome cé thé 1a monome (chi c6 mt phan ti protein) hoac polyme (cé trén 6 phan tit protein). Capsit c6 kha nang chiu nhiét, pH va cdc yéu t6 ngoai canh nén cé chic nang bao vé 16i axit nucleic. Trén mat capsit chifa cde thu thé dac higu gidp cho virut bam vao bé mat té bao. Capsit cing con chifa céc khang nguyén (KN) kich thich hé thong mién dich trong sudt thdi gian mac bénh. Phic hop bao gém capsit va axit nucleic duge goi 1A nucleocapsit. V6 capsit c6 kich thuée va cdch sip xép khac nhau khién cho cdc virut co hinh dang khac nhau. C6 thé chia ra ba loai céu trie : déi xting xoan, déi xing hinh khdi va cau tric phic tap (hinh 1.7; 1.8 ; 1.9). Capsome V6 ngoa lipoprotein ‘axit nucleic Capsit v6 protein Nucleocapsit Hinh 1.7. Cau trac cd ban cd virion. Léi la axit nucleic, vé capsit la protein. Léi va vé két hgp vi nhau goi la nucleocapsit. Phia ngoai capsit la vé ngoai lipoprotein V6i cdc gai glycoprotein. 15A B ic Hinh 1.8. A — So dé virut hinh que vdi cau tric di xing xan (virut Kham thudc 14) Capsome sp xép theo chiéu xoan cila axit nucleic. B - So dé virut da dign don gidn nhét. Mai mat la mét tam giae déu. Binh do 5 canh hop lai. Méi canh chtfa 3 capsome C ~ Su d6i xiing cia hinh da dién thé hién khi quay theo truc bac 2 (180°), bac 3 (120°) va bac 5 (72°), i 15) FA Hinh 1.9. Hinh thai ca mét sé loai virut. 1. Béi xing da dién A - polio, adeno, rota, B - Herpes ; 2. Odi xing xoain C - kham thude 14, D — cum, E ~ sgi, quai bi, parain fluenza, F - dai ; 3. 84i xing hn hgp G - virut pox, H - phage T chan. (Michael J. Pelczar et al. Microbiology, Concepts and Applications, 1993) 1.2.2.1. Cdu tric déi xttng xodn Bao gém céc virut gay bénh kham thuéc 14 (TMV), dai (rhabdo), quai bi, cim, séi (paramyxo, orthomyxo),... S6 di ¢6 cau tric xoin 1a do axit nucleic va protein sp xép theo kiéu xoan 1d xo xung quanh mot truc. Tuy loai ma 6 chiéu dai, dudng kinh va chu ky lap lai cita nucleocapsit khéc nhau, Céu tric xoan thuéng lam cho virut c6 dang hinh que hay hinh s 1612.2.2. Cdu tric déi xting hinh khéi 20 mat Khi quan sat k¥ céc virut “hinh cdu" duéi kinh hién vi dign tir ta thay thyc ra ching khong phai [a hinh cdu ma 1a hinh khdi da dién voi 20 mat tam gide déu, 30 canh va 12 dinh. Méi dinh 1a noi gap nhau cia 5 canh. Thude loai cau tic nay gém cdc virut adeno, reo, irido, herpes va picorna, Cch d6i xtmg theo 3 bac : bac 2, bac 3 va bac 5 (hinh 1.8c). Ta hay tudng tuong khi xoay 180° (bac 2), 120 °(bac 3) va 72° (bac 5) thi virut lai tr lai vi wri ci. Céc virut khdc nhau cé sé Iugng capsome trén méi canh khéc nhau do vay ma cé kich thudc khac nhau. $6 lugng capsome cing nhiéu thi virut co kich thudc cang In, Dya vao sé lugng capsome trén méi canh cé thé tinh duge téng sé capsome cia vo capsit theo cong thiic sau: N= 10 (n- 1)? + 2 trong dé N — téng sé capsome ctia capsit, n — sé capsome trén méi canh CAu tric kh6i thudng Jam cho virut c6 dang hinh cau Céu trie hén hgp Hay lay thé thuc khudn (TTK) lam vi du. TTK c6 cau tao dac biét, c6 dau phinh to kéo theo mot dudi dai, dac bigt phat trién & TIK — T chan (T2, T4, T6), con TTK - Te (13, T7) lai e6 dudi ngan hoac mot so tham chi khong ¢6 duéi. TTK — T2 c6 hinh tinh tring, dau cé cau tric dang khoi déi xuing voi 20 mat tam gide déu, néi voi dudi cé cdu utc déi xing xoan, nén ching la dang hén hop. 1.2.3, V6 ngoai Mét sé virut c6 vd ngoai (envelope) bao boc vd capsit. VS ngoai cé nguén goc tir mang sinh chat cia t€ bao duge virut cudn theo khi nay chéi. Vo ngoai cé edu tao gom 2 16p lipit va protein ~ Lipit gém photpholipit va glycolipit, hau hét bét ngudn tir mang sinh chat cha t€ bao (trit virut pox tir mang Golgi) vdi chtic naing chinh 1a én dinh cau tric cia virut. - Protein v6 ngoai thudng 1 glycoprotein cling c6 ngudén géc tir mang sinh chat. Tuy nhién, cing cé cée glycoprotein do virut m& hod duoc gan vao cdc vi ui chuyén biét trén mang sinh chat cia té bao, réi vé sau tré thanh cau tric bé mat cha virut. Vi du, cac gai hay cdc phan tir gp120 cia HIV hay hemaglutinin cia virut cdm, ching tuong tdc véi receptor cha té bio dé mo dau cho su xam nhap cia virut vao té bao. V6 ngoai cing cé thé cé nguén géc tir mang nhan (virut herpes). Va ngoai d bién tinh do tac dong cita cdc yéu t6 ngoai canh (dung méi hifu co, enzym...) lam mat kha nang gay nhiém cia virut. 1 Tigp theo sif tao thanh mARN cia virut [i téng hop protein. Virut can céc loai protein & céc thai diém khdc nhau vdi ham lugng khac nhau. Tuy thdt gian bigu hién ma protein cia virut duge chia thinh 3 loai : . Protein cia virut Fein be 17— Protein duge téng hop ngay sau khi nhiém goi la protein sdi tic protein khong edit trie thudng 1a cde enzym phuc vu cho sao chép axit nucleic. ~ Protein ditoe téng hyp mugn hon goi la protein mudn, thudng 1a protein cdiu tric tao nén v6 virut hod protein nim trong 1di. — Protein phan gidi thudng 18 enzym gitip giai phéng virut ra khdi té bao. Nhin chung, cd thdi gian xuat hién Lin sé lugng cla 3 loai protein nay déu dugc digu hoa sao cho hop ly nh& c6 céc gen diéu hoa am nhiém vu dong, md gen. Protein som 1a enzym xtc téc nen chi can mt lugng nhé, cdn protein mun 1a thanh phan cdu (ric nén bat budc phai tong hop véi sé lvgng Ién hon nhidu. Protein phan giai dugc téng hgp sau nita béi vi néu téng hop sém thi té bao chil sé bi tan trude khi mot lugng Ién virut dugc hinh thanh. Su nhiém virut gay xdo tron co ché diéu hod cia té bao. Sy tao thinh qua mitc cho phép axit nucleic va protein cha virut da lam suy sup hodn toan su téng hop cdc cao phan tir cia té bho. Cing cd tuéng hyp dién ra sy canh tranh gay gat gidta virut va 16 bao chil. Tuy nhién, trong ca hai truéng hgp tén sy diéu hod téng hgp cc thanh phan ctia virut duge kiém soat béi cdc gen cia virut hon 1 ciia té bao chi. Cac enzym cua virut Jogi virut cé chia cic enzym dic trumg cia minh, Cé thé chia ra Tam hai loai : ic enzym tham gia vao qué trinh sao ma va phién ma. - Céc enzym gitip virut xam nhap vao t€ bao va dua thé hé virut moi ra khdi t€ bao. Cac enzym cdn due phan thinh enzym cua virut va enzym cdm ting. Loai déu do virut mang theo, vi du enzym phién md nguge. loai sau duce téng hop nho genom cia virut va khong cé trong thanh phan cla virut truéng thanh. G virut orthomyxo (ciim) va paramyxo (4 clim) c6 enzym neuraminidaza khu tri & v6 ngoai gidp virut bam vio té bao. Polymeraza cia virut 18 enzym thyc hign sao chép. Néu polymeraza do té bio téng hgp dam ahigm duge chute nang doc ma axit nucleic ciia virut thi khéng cén thiét phai dua vio thanh phan cia virut méi, nguge lai néu khong dim nhiém duge chic nang doc ma thi nhat thiét phai dua né vao thanh phan cia virut thé he sau. ~ © virut ADN, genom luén duge ADN - polymeraza cia té bao chil dge ma, do vay virut van cé kha nang gay nhiém ma khong can mang theo polymeraza. - Virut chia ARN sgi (—) c6 bo gen agvoc (b6 sung) voi MARN. Muén nhan lén ching phi phien ma nh ARN ~ polymeraza phy thugc ARN cua riéng minh dé bién soi (-) thanh sgi (+) tue sqi MARN. Virut rerro mang theo enzym phién md nguoe dé chuyén hod ARN thanh ADN (vi cdc (@ bao dong vat khong cé enzym thye hién chit nang nay). 18~ Proteaza, Mot sé virut chita proteaza ding dé phan cat polyprotein, loai protein tién chat c6 kich thude 16n duoc hinh thanh sau dich mi tao ra ede protein chite nang 6 kich thude nhé hon. + Protein vo capsit cla virut picorna cé hoat tinh proteaza dic higu céin cho vige phan gidi protein trong nucleocapsit, gitip giai phong axit nucleic khdi v6 capsit. + MOt sé virut chtta endonucleaza va ligaza. 1.2.5. Tén goi va phan loai virut Cho dén nay, chua c6 mot hé théng phan loai virut nio ditge thye su chap nhan béi vin con thiu cdc dit ligu vé méi quan hé gitta céc virut va do dé virut cing khong cé 1én goi theo diing quy céch nhw 6 cdc vi sinh vat khde. Céch dat tén virut kha wy tién, chang han virut sdi (do gay bénh sdi), virut adeno gay bénh duting h6 hap trén Lin du tién phan lap duge tir hach hanh nhan (adenoid), virut coxsackie (do phan lap lin déu tign & Coxsackie — New york) hay virut Epstein — Barr (1a 1¢n hai nba nghién ctu phat hign ra chiing). Mot trong nhéng cach phan loai trude day 18 dya vao vat chi ma virut gay nhiém. Vi du virut ta lon, virwt otim Jon, virut dau bd, virut kham thude fd... Hé théng khée lai dya vao céc mo ma virut téc dong. Vi du virut tic dong vao hé théng ho hap thi goi Pneumotropic, vio hé than kinh goi 1a Neurotrepic, vao da goi 14 Dermatotropic, vao ndi tang goi la Viscerorropic. Céch phan loai nay tign loi cho gidi bac si va cée nha dich t@ hoc. Tuy nhién, nhd céc kign thiic vé cde dac diém Ly, hod, sinh hoc eta virut ngdy cang duoc tch lug ma Uy ban Quéc té vé phan joai virut goi tit Ia ICTV da dé nghi phan loai dya trén nhigu dac diém cau triic va sinh hoc cg ban. Uy ban nay léy don vi ho (family) lam trong. Céc chi tigu quy dinh ho bao gém : ~ Kiéu axit nucleic : ADN hay ARN. ~ Kiéu doi xing ca capsit : xan hay khéi hoac phitc tap. ~ Cé hay khong c6 vo ngoai. ~ Vint rap vO capsit véi axit nucleic — Vi tri gn v6 ngoai (mang té bao hay ming nhan). ~ Phan img voi ete va cac dung moi hoa tan Hipit khée. —36 lugng capsome. - Dudng kinh hat virut. — Kich thusc axit nucleic, Uy ban nay con théng hat ding tiép vi ngit - viridae dé chi tén ho, tiép vi ng - virinae dé chi ho phu, — virus dé chi chi. Tiép dau ngit cho méi ho thudng bat nguén tir 19dac diém dac trung cho ho. Vi du, Picornaviridae la ho virut ARN c6 kich thude nhd (pico = nhd, + ARN), Hepadnaviridae 1a virut ADN gay bénh gan (hepar = gan, + ADN). Retroviridae la virut ARN 6 enzym phién ma nguogc (retro = nguge). Mot sé ho khac mang ten do nguén géc lich sit. Céc quy dinh vé phan loai c6 dién khong duoc 4p dung cho virut. Dé xép cdc virut vao timg nhém lai phai ding them céc dic diém nhu lam sang, mién dich hoc va sinh hoc phan tit. Viée so sénh trinh ty axit nucleic cing duoc ding nhiéu dé xéc dinh m6i quan hé gitta céc virut. Vi tri trén ho 1a bd, c6 tip vi ngit 1a — virales. Dén nay méi duge xdc dinh chi c6 hai bO virut lién quan dén benh & ngudi 1a bo Mononegavirales (céc hg Filoviridae, Paramyxoviridae va Rhabdoviridae) va bO Nidovirales (Coronaviridae va Arteriviridae). Nam 1995, ICTV da cong bé hé thong phan loai téng hgp bao gém 1 bd, 71 ho, 11 ho phu, 164 chi va hon 4000 loai trong 46 cdn cé hang trim loai chua xc dinh do thiéu 36 liu. Céc virut méi cing khong ngimg duge phat hien. Hinh 1.10a va b gidi thiéu mot 86 virut thudng gap & dong vat va thuc vat. Khéng v6 ngoai C6 vd ngoai ds ADN ds ADN mauxenh adeno 2 S nguei Unha & Tho Poxiviridae Herpesviridae (vaccinia) (therpes simplex) ds ARN ssARN (reo type 1) ssARN SSADN Vigm xoang _ Retroviridae miéng (Rous sarcoma) 8 8 8 ? Ngoaiban —Polip 1 Uxo © éLgn Sngudi S chudt 400nm arenaviridae Coronaviridaé Bunyaviridae Togaviridae (ymphocytic —_(Viém ph€ (Bunyamweral) (Sindbis) choriomeningitis quan chim) Hinh 1.10a. Ho virut déng vat. Dudi tén ho la t&n virut dai dién (Michael J. Pelczar et al. Microbiology, Concepts and Applications, 1993) 20Khéng v6 ngoai C6 v6 ngoai dsADN ssARN ssARN g 0000 000 SS : Havirus Hordvirus (Visut kham soc dai mach) Nom virut Kham (visu soe thu la) Caulimovirus ae (Vint démhoas)! @ OB 8e6 Tobavirus Bromovius — Cucumovirus. == = {Virut kham dita) (Virut kh dua chuét ) dsARN ee ‘Tobamovirus (Virut kham thuéc 1a) Neptovirus —— (Nhe virut khaim du) Potexvirus (KKhoai tay X) ; dém vang 8 ry hoai tu Dianthovirus Cartavirus (Virut gay bénh em chusén Reoviridae | (Virut gay ddim khuyén cm chuéng) Sait eile “ @ Se ee SSADN _| nném veut ty hog tithus, Potyvirus (Khoai tay Y) gay lin ngs) = @ Geminivirus o Nhém virut gay 400nm | Luteovirus |d6m, héo ca chu (ina vang, cdl ope cf cai) Closterovirus (Virut vang cd cai) Hinh 1.10b. Ho va nhém virut gay bénh thyc vat. Dudi ten nhém hay ho 1a tén virut dai dién. (Michael J. Pelczar et al. Microbiology, Concepts and Applications, 1993) 1.2.5.1. Mé6t 86 dée diém cua virut dang trong phan loai dén lodi Dac diém ciia virion —Hinh théi + Kich thuéc + Hinh dang + C6 hay khong cé peplome, ban chat ciia peplome + C6 hay khong c6 vd ngoai + Cu tric va kiéu déi xting cia capsit —Tinh chat vat I§ va ly hod + Kh6i lugng phan tir (Mr) + Mat do néi (buoyant density) cia virion (trong CsCl, Sa +) + Hé s6 ling cia virion 21+ Do bén pH + Do bén nhiet + BO bén cation (Mge"*, Mn?*) + Do bén voi dung moi + D6 bén véi chat tay rita + Dé bén phong xa —Genom + Loai axit nucleic (ADN hay ARN) + Kich thuéc genom, tinh bing cap nucleotit (bp) vai mach kép hoac bing sé nucteotit (b) véi mach don. + S6 mach - mach don hay kép + Dang thang hay khép vong + Phan cuc — soi am hay duong + 86 doan va kich thuéc cic doan + Trat ty nucleotit + Surcé mat cla doan lap lai + Su cé mat cla isomer hod +TyieG-C + Cé hay khong cé mii d dau $* + C6 hay khéng cé protein lign két & dau 5" + C6 hay khong cé dudi poly A 6 dau 3° — Protein + S6 luong, kich thudc, chitfc nang cilia protein cu tric. + $6 luong, kich thudc, chiéc nang cia protein khong cdu tric (Ia cdc enzym). + Chi tiét vé chide nang chuyén biét cua protein dac biét 1a transcriptaza, enzym phién ma ngugc, hemaglutinin, neuraminidaza va hoat tinh dung hop. + Tribh ty axit amin + Su gan duing vio protein (glycosyl hod) va gdn photpho vao protein (photphoryl hoa). + Lap ban dé epitop — Lipit + Him lung, dac tinh,... -Té chite genom va sao chép +Té chie gen + Dac diém sao ma + $6 lugng va vi tri khung doc mé (ORF)+ Dac diém phién ma + Noi tich lu protein cla virion + Noi lip rap + Noi virion truéng thanh va phéng thich —Tinh chat KN + Méi quan hé vé huyét thanh hoc — Dac diém sinh hoe + Loai vat cho ty nhién + Phuong thie lan truyén trong ty nhién + M6i quan hé cia vecto + Phan bé dia ly + Kha nang gay bénh + Tinh huéng m6, gay bénh ly md. 1.2.5.2 Phan logi virut thuc vat n phan loai virut thyc vat goi tat 1A PSV (Plant virus subcommittee) quy dinh — V-virus — Vd-viroid -M ~ kham (mosaic). Vi dy virut kham dau dila vict tat 1a CPMV (Cowpea mosaic virus) — Mo — dm (mottl). Vi du virut dém dau diia viet tat lA CPMoV (Cowpea mottle virus) — RS — dém vong (ring spot). Vi dy virut dém vong thude 14 viet tat la TRSV (Tobacco ring spot virus) — Mot s6 truéng hgp dé khéi nhaim lan tén cay duge viét bing 2 chit cdi nhung chit sau viet nhé hon chit trudc. Vi du virut dém vang cd chua viet tit la TORSV (Tomato ring spot virus) — Virut lign quan dén mot s6 bénh cha cay thi thém chit a (associated) tneéc chit V. Vi du virut soc nau san viét tat Li CBSaV (cassava brown steak associated virus). Virut thye vat duge xép vao cdc nhém dua theo cde dac diém hinh théi cia capsit, s6 lugng va két ciiu axit nucleic. Virut thuc vat gém 14 ho, 70 chi trong dé cé 27 chi chua dugc xép vio ho nao, ngoai ra cdn nhiéu Loai chua duge xép vio chi nto. C6 hai loai Lodi : — Loai da xde dinh (definitive species) — Loai méi chi 1a gid dinh (putative species). Cc ho virut thyc vat duoc xép vio 5 nhom sau day : 1. ADN chuéi kép — Ho Caitimaviridae 232. ADN chuéi don - Ho Circoviridae - Ho Geminiviridae 3. ARN chudi kép ~ Ho Partitiviridae ~ Ho Reoviridae 4. ARN chudi don, am B6 Mononegaviridae - Ho Rhabdoviridae ~ Ho Bungaviridae 5. ARN chudi don, duong ~ He Bromoviridae 1.2.5.3. Phén logi thé thuc khuén Cach goi ten TTK citng khong theo quy luat, chi yéu IA phu thuéc vio nha nghién ctu. TTK duge chia ra cdc ho sau day : 1.2.5.3.1. Khéng 06 vé ngodi a) Chita ADN chudi kép - Hg Myoviridae. TTK dau hinh kh6i. Dai dién P2. TTK dau hinh koi dai. Dai dién T2. — Ho Siphoviridae. Dai dien 2. — Ho Podoviridae. Dai dién T7 (dudi ngin). ~ Ho Corticoviridae. Dai dién PM2 (khong dudi). ~ Ho Tectiviridae. Dai dién PRDI (khong dudi), Bam 35. b} Chita ADN chudi don ~ Ho Mnoviridae. Dai dién MV - L1 (dang que), phage 4, fd (dang soi) ~ Ho Microviridae. Dai dién @X 174 (dang khoi, khong dudi) ¢) Chita ARN chudi dan ~ Ho Leviridae. Dai dién MS2 (dang khdi, khong dudi), R17 1.2.5.3.2. C6 ud ngodi a) Chita ADN chudi kép ~ Ho Plasmaviridae. Dai dign MV ~ L2 (hin cau) 5) Chita ARN chudi kép ~ Ho Cystoviridae. Dai dign 6 (dang khéi) 1.2.5.4, Phan logi virut dua vao vét chit Dua vao vat chil, Prescott di chia virut thanh 6 nhém Idn. (Prescott et al, Microbiology Mc Graw Hill, 2002). 241.2.5.4.1. Virut vita nhén lén é d6ng vdt cé xwong séng vita nhén lén é céc co thé khée Nhiéu thinh vién cita cdc ho Reoviridae, Bunyaviridae, Rhabdoviridae va Togaviridae vita c6 thé nhan lén 6 DVCXS vita cé thé nhan lén & cdc co thé khiic. Mot 86 ho khdc cé cae chi chi cé thé nhan lén 6 DVCXS. a) Ho Iridoviridae Cac hat hinh da di¢n (125 — 300nm) bao gom nucleocapsit hinh cdu duge bao béi 1ép lipit cd bé sung cac tiéu phan protein. Cé thé 6 VO ngoai nhung vé nay khong cin cho qué trinh gay nhigm. ADN kép (khdi long phan tir 100 - 200 x 10°), khép kin véi doan Jap lai cing chiéu @ cudi phan tit, chtta mot sé enzym. Phién ma va téng hop ADN trong nhan, mARN khong cé dudi poly A, dp rép trong sinh chat. Céc chi dai dién : ~Iridovirus (virut sac edu véng kich thudc nhd — 120nm ¢ con tring). —Chlorviridovirus (virut séc céu véng kich thuée 1én — 180am ¢ c6n tring). ~ Ranavirus (nhém virut & &ch nhdi) — Lymphocystivirus (virut 8 c4) b) Ho Poxviridae ADN kép (trong lung phan tir 85 — 240 x 10°) vdi doan lap lai nguoc chiéu d cuéi phan tir. La virut c6 kich thuée 1én nhat (170 — 260 x 300 - 450nm). Cau tric phite tap g6m nhiéu 1ép, trong 46 66 16p lipit. Loi chita tat cd cdc enzym can cho téng hyp mARN, nhan lén trong sinh chat. Cac chi dai die — Orthopoxvirus (virut vaccinia va cdc virut c6 quan hé he hang). ~ Molluscipoxvirus (virut cia ngudi nhiém qua nhuyén thé). —Avipoxvirus (virut pox 6 gia cam va cdc virut ho hang). — Parapoxvirus (milkpox va cdc virut ho hang) gém 4 chi trén nhiém 6 DVCXS. — Entomepoxvirus (virut pox & con wing). ¢) Ho Parvoviridae Hat hinh da dién kich thuéc 18 - 22am, ADN don (Khoi lugng phan tir 1,5 — 2,0 x 10°). Khong chia enzym. Lap rap trong nhan. 2s