ç•ªèŒ„æ–‘èŽç— æ¯’å±¬ç— æ¯’ä¹‹ç™¼ç”Ÿèˆ‡å¿«é€Ÿåµæ¸¬æŠ€è¡“ä¹‹æ–°ç™¼å±•

8:125-132, 1999s d y e h @ n c h u . e d u . t w 04 - 2852501881 03 0. 1999. . 8 : 125 - 132.(the genus To s p o v i ru s) B u n y a v i r i d a eTo s p o v i ru s(Thrips palmi Karny) (Wa t e r m e l o nsilver mottle viru s, WSMV) To s p o v i ru s(S c i rtothrips dorsalis Hood) (Peanut chlorotic fan-spot viru s, PCFV)S RNA(13,TSWV 16) 1915(spotted wilt) B u n y a v i r i d a e(arthropods) ( 3 ) 1 930To s p o v i ru sB u n y a v i r i d a e(Tomato spotted wilt viru s, TSWV) ( 37 ) ( 36 ) T S W V( 33 ) (larval stages) 828 00( 32 ) ( 31 ) (20, 27, 29, 33) ( 44 ( s p o t )) ( c h l o r o s i s ) ( m o t t l i n g ) ( s t u n t i n g ) 80(wilting) ( n e c r o s i s )110 nm( 42 ) ( l i p o p r o t e i ne n v e l o p e )R N A1970L RNA (9 kb)M RNA (5 kb) S(tomato spotted wilt virus RNA (3-3.5 kb) ( 17 ) R N A g r o u p ) 1991( I n t e r n a t i o n a l (nucleocapsid protein, NP) Committee on Taxonomy of Virus, ICTV) (pseudo-circular structure) ( 28 ) To s p o v i ru s B u n y a v i r i d a e(negative sense and

1 2 68 4 1999ambisense coding strategies) ( 17 ) R N A (N protein)G 1 (ELISA) ( w e s t e r nG 2 (G1/G2 glycoproteins) R N A ( L blotting) (monoclonal antibody) p r o t e i n )N S mN S s (NSm and NSsp r o t e i n s )L RNATo s p o v i ru s (L protein)MS RNAs199510, 19, 45,48) T S W V(movement protein) ( 23 ) G1/G2 (serogroup I) G 1G 2( s p i k e s )S RNAN S s(G roundnut ringspot viru s, GRSV)(Tomato chlorotic spot viru s, TCSV) ( 22 ) ( N P )T S W V( 977) (Impatiens necrotic spot viru s, INSV) 1990 L a wT S W VG 1G 2 ( 48 (Watermelon silver mottle viru s, WSMV) ) Prins and Goldbach (Peanut bud necrosis viru s,1998 (32) P B N V ) ( 39 ) R N A19985Wa g e n i n g e n R N ATo s p o v i ru s1 3 ( s p e c i e s ) WBNV (Watermelon bud necrosis viru s)(21)m R N AW S M VP B N VPYSV (Peanut yellow spot viru s) ( 38 ) IYSV (I r i sR N Ayellow spot viru s) ( 7 ) PSMV (Physalis severe mottle viru s)( 8) CSNV (C h rysanthemum stem necro s i s v i ru s) ( 11 ) Z L C V( 22 N S s) (Zucchini lethal chlorosis viru s) ( 11 ) PCFV (P e a n u tc h l o rotic fan-spot viru s) (50) (rough endoplasmia reticulum, RER) ( ) (budding) L ( ) R N AN S m(23, 41) R N AT S W VI N S VW S M VP B N VLMS RNAsT S W VL RNA8 897( 15 ) M RNA N S mR N ATo s p o v i ru s( 2 5 ) (open reading frame, ORF)(negative polarity)331.5 kDaRNA I N S VL RNA 8876( 4 3 ) 330.3 kDa T S W V69 . 5P B N VLR N A8 911330 kDa24405 0 W S M VL RNA 8917331.8 kDa( 20 ) P B N V 91 . 3( 1 ,( 1 8 )

127Table 1. Amino-acid sequence identities (%) of the N genes of tospoviruses.S e r o g r o u p I I I I I I I V V V I V I I V I I I I X XVi r u s T S W V T C S V G R S V I N S V W S M V P B N V W B N V P Y S V I Y S V P S M V C S N V Z L C V P C F VT S W V 100 77 78 55 33 33 32 26 34 29 77 73 19T C S V 100 81 55 26 33 31 23 34 29 75 73 22G R S V 100 54 33 34 33 22 33 29 75 75 23I N S V 100 30 30 29 23 30 28 58 52 22W S M V 100 86 85 24 44 57 33 32 22P B N V 100 85 23 44 57 32 31 24W B N V 100 22 43 58 32 30 21P Y S V 100 21 23 25 20 66I Y S V 100 49 35 32 27P S M V 100 34 29 22C S N V 100 82 22Z L C V 100 21P C F V 100Data from the Fourth International Symposium on Tospoviruses and Thrips in Floral and Vegetable Crops (Wageningen, TheNetherlands, May 2-6, 1998, communicated by Dr. R. Kormelink). TSWV: To m a t o spotted wilt viru s (Australia, USA, TheNetherlands), TCSV: Tomato chlorotic spot viru s (Brazil), GRSV: G roundnut ringspot viru s (South Africa), INSV: I m p a t i e n sn e c rotic spot viru s (USA, Europe), WSMV: Watermelon silver mottle viru s ( Taiwan, Japan), PBNV: Peanut bud necrosis viru s(India), WBNV: Watermelon bud necrosis viru s (India), IYSV: Iris yellow spot viru s (The Netherlands), PSMV: Physalis severemottle viru s ( Taiwan), CNSV: C h rysanthemum stem necrosis viru s (Brazil), ZLCV: Zucchini lethal chlorosis viru s ( B r a z i l ) ,P C F V: Peanut chlorotic fan-spot viru s ( Ta i w a n )T S W VI N S V 44 . 3 46 . 5G R S VT C S V() T S W VI N S VW S M VP B N VM RNAT S W VT S W V4 821( 24 ) I N S V4 972( 2 6 ) W S M V4 8 8 0 ( 6 ) P B N V4 8 0 1 ( 4 0 ) M RNA(ambisense coding strategy)I N S V6 3 . l 55 . 3O R F s (viral sense strand) RNAde Av i l a G R S VT C S V 94 . 4 81 . 534 kDa(NSm) (TSWV:33.6 kDa, INSV: 34 . 1kDa, WSMV:35 kDa, PBNV:34 kDa)( v i r a l G R S VI N S V Scomplementary strand) 130 kDa R N A2 9925' ( T S W V:127.4 kDa, INSV:124.9 kDa, WSMV:127.6 kDa,P B N V:127.6 kDa) G 1G2 ( 14 ) W S M V 3534To s p o v i ru sM RNAB u n y a v i r i d a eR N A275439B u n y a v i r i d a e M N S s WSMV S RNAR N AT S W VI N S VG 1 / G 2 T S W VI N S V 6 l 85 42( 35 R G D) W S M VS RNA1261W S M VP B N VM RNA G l / G 2T S W VINSV 7526 13( 51 ) P B N VS(6,40) ( 93' ) 1 993P a n gT S W V- B S RNA3 049T S W V 77 . 579 . l( 30 ) G R S V T S W V 67N S s 51 . 9R N A3 057( 39 S RNATo s p o v i ru s49.5 kDaN S s 30.6 kDa) 13W S M V8 6W B N V( )T S W VS RNA 291627513 R N A29W S M VP B N V7 9R N A 52.4 kDa(NSs protein)8 1444 6( 13 R N A 28.8 kDa) ( 21 ) W S M VP B N VW B N V

1 2 88 4 1999P Y S V (49)S RNA 2970R N A53.2 kDaN S sR N A(C u c u r b i t a c e a e)28.0 kDaN222 6(Thrips palmi K a r n y ) P Y S VS RNAIT S W V I I7.5 kDaG R S VT C S V I I I ( 38 ) I Y S VS RNA3 105( 7 ) I N S V50.1 kDaNSs protein30.5 kDa( 46 44W S M V) W S M VP S M VI Y S V( To s p o - To ) ( 2 ) ( 20 W S M V) 58C S N VZ L C VP B N V (34) W B N V (21) 8 27577P C F V P Y S V 661 92 7 SM RNAA - UW S M VS RNAO R F35345' 3' 225 751 ( p a n h a n d l e) M RNA 4880N S mG 1 / G 2s t r u c t u r e ) R N A( p s e u d o - c i r c u l a rs t r u c t u r e )R N A 8917T S W V( 17 ) I N S V224 7 ( u n p u b l i s h e dS RNA3' 8d a t a )3' 5 '- A U U G C U C U - 3 '( 5 '- A U U G C U C U - 3 ')LMS R N AT S W VI N S VW S M VP B N VLW S M VTo s p o v i ru sM 3' W S M V( 1 2 ) To s p o v i ru sB u n y a v i r i d a eR N A3' 8( 2 9 ) I V ( 46 ) To s p o v i ru s( 6T S W VI N S V 314 1) L 1992(49)W S M V 1988(S c i rtothrips dorsalis H o o d )1990(Peanut chloro t i cfan-spot viru s, PCFV) ( 5 ) I(T h r i p s p a l m i T S W V I IKarny) G R S V I I IT C S V I ( 48 W S M VW S M V) 1998To s p o v i ru sS RNA( 49 ) 2833W S M V5' 3' (NSs) 51 . 5( 4) k D a19 . 35 4 . 2 P C F V31.1 kDa( 4 7 ,

12922 . 36 7 . 53' 5'-AUUGCUCU-3' (unpublished data)P C F VTo s p o v i ru s 13W S M VP C F VPhysalis severe mottle( 8v i ru s (PSMV) ) T S W VG R S I N S VWSMV P C F VP C F VII VIT S W V I IS G R S VR N AP C F V T S W VI N S VW S M VPCFV L RNA1 . Adam, G., Lesemann, D. E., and Vettern, H. J. 1991.( d e g e n e r a t e Monoclonal antibodies against tomato spotted wilt virus:p r i m e r s )T S W V characterization and application. Ann. Appl. Biol. 118 : 87 -G R S VI N S V 104 .W S M VP C F V2 . Adam, G., Yeh, S. D., Reddy, D. V. R., and Green, S. K.1993. Serological comparison of tospovirus isolates fromTe n u i v i ru s Taiwan and India with impatiens necrotic spot virus andTo b a m o v i ru sP o t e x v i ru sP o t y v i ru sN e p o v i ru s serogroup of tomato spotted wilt virus. Arch. Vi r o l .C u c u m o v i ru s130 : 237 - 250 .3 . Brittlebank, C. C. 1919. Tomato disease. J. Agric.Victioria. 17:213-235.4 . Chen, C. C., Ho, H. M., Chang, T. F., Chao, C. H., andT S W VG R S VI N S Yeh, S. D. 1995. Characterization of a tospovirus-like W S M Vvirus isolated from wax gourd. Plant Prot. Bull. 37:117 -131 .5 . Chen, C. C., and Chiu, R. J. 1996. A tospovirus infecting

1 3 08 4 1999peanut in Taiwan. Acta Horticult. 431:57-67.6 . Chu, F. H., and Yeh, S. D. 1998. Comparison ofreplication forms and ambisense M RNA of watermelonsilver mottle virus with other tospoviruses.Phytopathology 88:351-358.7 . Cortes, I., Livieratos, I. C., Derks, A., Peters, D., andKormelink, R. 1998. Molecular and serologicalcharacterization of iris yellow spot virus, a new anddistinct tospovirus species. Phytopathology 88:1276-1282 .8 . Cortes, I., Pereira, A., Goldbach, R., Peters, D., andKormelink, R. 1998. An RT-PCR procedure to amplify SRNA sequences of distinct tospoviruses. In: Recentprogress in Tospovirus and thrips research. Edited byPeters, D., and Goldbach, R., Agricultural University,Wageningen, The Netherlands. P35-37 (abstract).9 . de Avila, A. C., de Haan, P., Kormelink, R., de OliveiraResende, R., Goidbach, R., and Peters, D. 1993.Classification of tospovirus based on phylogeny ofnucleoprotein gene sequence. J. Gen. Virol. 74:153-159.10 . de Avila, A. C., Huguenot, C., de Oliveira Resende, R.,Kitakima, E. W., Goldbach, R., and Peters, D. 1990Serological differentiation of 20 isolates of tomato spottedwilt virus. J. Gen. Virol. 71:2801-2827.11 . de Avila, A., Pozzer, L., Bezerra, I., Kormwlink, R.,Prins, M., Peters, D., Nagata, T., Kitajima, E., andResende, R. 1998. Diversity of tospoviruses in Brazil. In:Recent progress in Tospovirus and thrips research. Editedby Peters, D., and Goldbach, R., Agricultural University,Wageningen, The Netherlands. P32-34 (abstract).12 . de Haan, P., Wagemakers, L., Peters, D., and Goldbach,R. 1989. Molecular cloning and terminal sequencedetermination of the S and M RNAs of tomato spottedwilt virus. J. Gen. Virol. 70:3469-3473.13 . de Haan, P., Wagemakers, L., Peters, D., and Goldbach,R. 1990. The S RNA segment of tomato spotted wilt virushas an ambisense character. J. Gen. Virol. 71:1001-1007.14 . de Haan, P., Avila, A. C., Kormelink, R., Westerbroek, A.,Gielen, J. J. L., Peters, D., and Goldbach, R. 1992. Thenucleotide sequence of the S RNA of impatiens necroticspot virus, a novel tospovirus. FEBS Letters 306:27-32.15 . de Haan, P., Kormelink, R., de O. Resende, R., vanPoelwijk, F., Peters, D., and Goldbach, R. 1991. To m a t ospotted wilt virus L RNA encodes a putative RNApolymerase. J. Gen. Virol. 71:2207-2216.16 . Francki, R. I. B., Fauquet, C. M., Knudson, D. L., andBrown, F. 1991. Classification and Nomenclature ofViruses. Fifth Report of the International Committee onTaxonomy of Viruses. Arch. Virol. Suppl. 2.17 . German, T. L., Ullman, D. E., and Moyer, J. W. 1992.Tospoviruses: diagnosis, molecular biology, phylogeny,and vector relationships. Ann. Rev. Phytopathol. 30:315-348 .18 . Gowda, S., Satyanarayana, T., Naidu, R. A., Mushegian,A., Dawson, W. O., and Reddy, D. V. R. 1998.Characterization of the large (L) RNA of peanut budnecrosis tospovirus. Arch. Virol. 143:2381-2390.19 . Gonsalves, D., and Trujillo, E. E. 1986. Tomato spottedwilt virus in papaya and detection of the virus by ELISA.Plant Dis. 70:501-506.20 . Iwaki, M., Honda, T., Hanada, K., Tochihara, H., Yo n a h a ,T., Hokama, K., and Yokoyama, T. 1984. Silver mottledisease of watermelon caused by tomato spotted wiltvirus. Plant Dis. 68:1006-1008.21 . Jain, R. K., Pappu, R. H., Pappu, S. S., Krishna Reddy,M., and Vani, A. 1998. Watermelon bud necrosistospovirus is a distinct virus species belonging toserogroup IV. Arch. Virol. 143:1637-1644.22 . Kormelink, R., Kitajima, B. W., de Haan, P., Zuidema, D.,Peters, D., and Goldbach, R. 1991. The nonstructuralprotein (NSs) encoded by the ambisense S RNA segmentof tomato spotted wilt virus is associated with fibrousstructures in infected plant cells. Virology 181:459-468.23 . Kormelink, R., Storms, M., van Lent, J., Peters, D., andGoldbach, R. 1994. Expression and subcellular locationof the NSm protein of tomato spotted wilt virus (TSWV),a putative viral movement protein. Virology 200:56-65.24 . Kormeniink, R., de Haan, P., Meurs, C., Peters, D., andGoldbach, R. 1992. The nucleotide sequence of the MRNA segment of tomato spotted wilt virus, a bunyaviruswith two ambisense RNA segment. J. Gen. Vi r o l .73 : 2795 - 2804 .25 . L a w, M. D., and Moyer, J. W. 1990. A tomato spotted wiltlike virus with a serologically distinct N protein. J. Gen.Virol. 71:933-938.26 . L a w, M. D., Speck, J., and Moyer, J. W. 1992. The MRNA of impatiens necrotic spot tospovirus (B u n y a v i r i d a e) has an ambisense genomic org a n i z a t i o n .Vi r o l o g y. 188:732-741.27 . Matthews, R. B. F. 1991. Plant virology (3rd ed.). 835pages, Academic Press, New Yo r k .28 . Mohamed, N. A. 1981. Isolation and characterization ofsubviral structures from tomato spotted wilt virus. J. Gen.Virol. 53:197-208.

13129 . M u r p h y, F. A., Fauquet, C. M., Bishop, D. H. L.,Ohabrual., S. A., Jarvis, A. W., Martelli, O. P., Mayo, M.A., and Summers, M.D. 1995. Classification andNomenclature of Viruses. Sixth report of the internationalcommittee on taxonomy of viruses. Arch. Virol. Suppl.10 .30 . Pang, S. Z., Slightom, J. L., and Gonsalves, D. 1993. Thebiological properties of a distinct tospovirus and sequenceanalysis of its S RNA. Phytopathology 83:728-733.31 . Peters, D., Wijkamp, L., van de Wetering, F., andGoldbach, R. 1996. Vector relations in the transmissionand epidemology of tospoviruses. Acta. Horticult. 431:29-42 .32 . Prins, M., and Goldbach, R. 1998. The emerging problemof tospovirus infection and nonconventional methods ofcontrol. Tren. Microbiol. 6:31-35.33 . R e d d y, D. V. R., and Wightman, J. A. 1988. To m a t ospotted wilt virus: Thrips transmission and control. Adv.Dis. Vector Res. 5:203-220.34 . R e d d y, D. V. R., Ratna, A. S., Sudarshana, M. R., Poul, F. ,and Kiran Kumar, I. 1992. Serological relationships andpurification of bud necrosis virus, a tospovirus occurringin peanut (Arachis hypogaea L.) in India. Ann. Appl.Biol. 120:279-286.35 . Ruoslahti, E. and Pierschbacher, M. D. 1986. Arg - G l y -Asp: A versatile cell recognition signal. Cell 44:517-518.36 . Sakimura., K. 1962. The present status of thrips-bornedisease, in Maramorocsh. K. (ed): Biological transmissionof disease agents. New York and London, AcademicPress, p33-40.37 . Samule, G., Bald, J. G., and Pittman, H. A. 1930.Investigations on "spotted wilt" of tomatoes. AustraliaCommonwealth Coun. Sci. Ind. Res. Org. Bull. 44:1-64.38 . Satyanarayana, T., Gowda, S., Lakshminarayana Reddy,K., Mitchell, S. E., Dawson, W. O., and Reddy, D. V. R.1998. Peanut yellow spot virus is a member of a newserogroup of To s p o v i ru s genus based on small (S) RNAsequence and organization. Arch. Virol. 143:353-364.39 . Satyanarayana, T., Mitchell, S. B., Reddy, D. V. R.,Brown, S., Kresovich, S., Jarret, R., Naidu, R. A., andDemski, J. W. 1996. Peanut bud necrosis tospovirus SRNA: Complete nucleotide sequence, genomeo rganization and homology to other tospoviruses. Arch.Virol. 141:85-98.40 . Satyanarayana, T., Mitchell, S. B., Reddy, D. V. R.,Kresovich, S., Jarret, R., Naidu, R. A., Gowda, S., andDemski, J. W. 1996. The complete nucleotide sequenceand genome organization of the M RNA segment ofpeanut bud necrosis tospovirus and comparison with othertospoviruses. J. Gen. Virol. 77:2347-2352.41 . Storms, M. M. H., Kormelink, R., Peter, D., Van Lent, J.W. M., and Goldbach, R. 1995. The nonstructural NSmprotein of tomato spotted wilt virus induces tubularstructures in plant and insect cells. Virology 214:485-493.42 . van Kammen, A., Henstra, S., and Le, T. S. 1966.Morphology of tomato spotted wilt virus. Vi r o l o g y30 : 574 - 577 .43 . van Poelwijk, F., Prins, M., and Glodbach, R. 1997.Completion of the impatiens necrotic spot virus genomesequence and genetic comparison of the L proteins withinthe family Bunyaviridae. J. Gen. Virol. 78:543-546.44 . Verkleij, F. N. and Peters, D. 1983. Characterization of adefective form of tomato spotted wilt virus. J. Gen. Vi r o l .64 : 677 - 686 .45 . Wang, M., and Gonsalves, D. 1990. ELISA detection ofvarious tomato spotted wilt virus isolates using specificantisera to structural proteins of the virus. Plant Dis.74 : 154 - 158 .46 . Yeh, S. D., Lin, Y. C., Cheng, Y. H., Jih, C. L., Chen, M.J., and Chen, C. C. 1992. Identification of tomato spottedwilt-like virus infecting watermelon in Taiwan. Plant Dis.76 : 835 - 840 .47 . Yeh, S. D., and Chang, T. F. 1995. Nucleotide sequence ofthe N gene of watermelon silver mottle virus, a proposednew member of the genus Tospovirus. Phytopathology85 : 58 - 64 .48 . Yeh, S. D., Chao, C. H., Cheng, Y. H., and Chen C. C.1996a. Serological comparison of four distinctTospoviruses by polyclonal antibodies to purifiednucleocapsid protein. Acta Horticult. 431:122-134.49 . Yeh, S. D., Cheng, Y. H. Jih, C. L., Chen, C. C., andChen, M. J. 1988. Identification of tomato spottedinfecting horn melon and watermelon. Plant Prot. Bull.30 : 319 - 420 .50 . Yeh, S. D., Peng, Y. C., Chao. C. H., and Chen, C. C.1998. Peanut chlorotic fan-spot virus is serologically andphylogenetically distinct from other tospoviruses. In:Recent progress in Tospovirus and thrips research. Editedby Peters, D. and Goldbach, R., Agricultural University,Wageningen, The Netherlands. P. 42-43 (abstract).51 . Yeh, S. D., Sun, I. J., Ho, H. M., and Chang T. F. 1996b.Molecular cloning and nucleotide sequence analysis ofthe S RNA of watermelon silver mottle virus. ActaHorticult. 31:244-260.