116 Penaeid shrimp (ton) Whiteleg shrimp (ton) ton Giant tiger shrimp (ton) Penaeid shrimp (USDx1000) USDx1000 5,000,000 30,000,000 4,500,000 4,000,00

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1 Fish Pathology, 52 (3), , The Japanese Society of Fish Pathology * White Spot Disease: WSD ( = Penaeid Acute Viremia: PAV) Mari Inada 1, Tohru Mekata 1 and Toshiaki Itami 2 * 1 National Research Institute of Aquaculture, Japan Fisheries Research and Education Agency, Mie , Japan 2 Faculty of Agriculture, University of Miyazaki, Miyazaki , Japan (Received July 27, 2017) ABSTRACTWhite spot disease (WSD), the synonymous of penaeid acute viremia (PAV), has become one of the most serious problems of the shrimp culture business in the world. One important infection route of the causative virus (WSSV: white spot syndrome virus = PRDV: penaeid rod-shaped DNA virus) is a vertical transmission from spawner to larvae via eggs in the seed production process of kuruma shrimp (Marsupenaeus japonicus). To prevent the vertical transmission, disinfection methods of fertilized eggs and high-sensitive detection methods for the causative virus by molecular technologies have been developed. Until recently, the seed production has been depending on the mated and captured female broodstock. However, as WSSV is spontaneously affecting the broodstock in nature at some level, the vertical infection occurs due to the virus replication in the virus-carrier female broodstock caused by spawning stress. Therefore, specific pathogen free (SPF) kuruma shrimp stock is highly important for prevention of WSD and for sustainable shrimp culture in Japan Fig. 1 white spot disease: WSD = penaeid acute viremia: PAV 1992 Chou et al., * Corresponding author itamit@cc.miyazaki-u.ac.jp Lo et al., g

116 Penaeid shrimp (ton) Whiteleg shrimp (ton) ton Giant tiger shrimp (ton) Penaeid shrimp (USDx1000) USDx1000 5,000,000 30,000,000 4,500,000 4,000,000 25,000,000 3,500,000 20,000,000 3,000,000

2 116 Penaeid shrimp (ton) Whiteleg shrimp (ton) ton Giant tiger shrimp (ton) Penaeid shrimp (USDx1000) USDx1000 5,000,000 30,000,000 4,500,000 4,000,000 25,000,000 3,500,000 20,000,000 3,000,000 2,500,000 15,000,000 2,000,000 1,500,000 10,000,000 1,000,000 5,000, , year Fig. 1.Annual shrimp production volume and values in the world. (from FishStat Plus) WSD Satoh, 2012 Momoyama and Muroga, RV-PJrod-shaped nuclear virus of Penaeus japonicus PAV PRDVpenaeid rod-shaped DNA virus OIE WSDwhite spot disease WSSVwhite spot syndrome virus Flegel2012 Thitamadee et al.2016 NimaviridaeWhispovirus Mayo, 2002; Vlak, nm nm nm nm Fig. 2A BVlak, kb DNA Van Hulten et al., 2001; Yang et al., VP664VP28 A B Fig. 2. (A) Transmission electron micrograph of the WSSVinfected primary ovarian cells of kuruma shrimp. Magnification : 5,000. By courtesy of Dr. M. Maeda. (B) High-magnification of the WSSV virions replicating in the infected shrimp ovarian cell nucleus. Magnification : 17,000. By courtesy of Dr. M. Maeda. VP26VP24VP19 VP15 Leu et al., 2005; Tsai et al., 2006; Xie et al., 2006GenBank 3 WSSV DNA AF369029

117 AF440570AF332093 80 Fig. 3A B 0.5 μm Momoyama and Muroga, 2005 PCR nested-pcr PCR LAMP Durand and A B Fig. 3. (A) Gross view of the WSSV-infected shrimp (upper) and normal shrimp (lower).

, 2004; Lo et al., 1996 in situ hybridizationdot blot hybridization dot immunoblot assay ELISA Maeda et al., 2004 PCR 5 mg/l 5 Satoh et al.

3 117 AF440570AF Fig. 3A B 0.5 μm Momoyama and Muroga, 2005 PCR nested-pcr PCR LAMP Durand and A B Fig. 3. (A) Gross view of the WSSV-infected shrimp (upper) and normal shrimp (lower). Arrows show the typical white spot on the carapace. (B) Arrows show the typical white spots on the removed carapace. Lightner, 2002; Kono et al., 2004; Lo et al., 1996 in situ hybridizationdot blot hybridization dot immunoblot assay ELISA Maeda et al., 2004 PCR 5 mg/l 5 Satoh et al., mg/l 18 L/min * 4 6 Mushiake et al., 1998 PCR 2014 PCR 2014 PCR * 25p. 139.

4 Muroga, 2001 Namikoshi et al., 2004; Witteveldt et al., 2004; Kono et al., 2014 VP28 DNA Kono et al., Penaeus monodon white leg shrimp; Litopenaeus vannamei EVE; endogenous viral elementsutari et al., 2017 Dang et al., 2010; Utari et al., 2017 Dang et al., * 1993 * 29p PCR PCR 1 WSSV SPF WSSV SPF L. vannamei acute hepatopancreatic necrosis disease: AHPND, Enterocytozoon hapatopenaei: EHP 28 Chou, H.-Y., C.-Y. Huang, C.-H. Wang, H.-C. Chiang and C.-F. Lo1995: Pathogenicity of a baculovirus infection causing white spot syndrome in cultured penaeid shrimp in Taiwan. Dis. Aquat. Organ., 23, Dang, L. T., T. Koyama, A. Shitara, H. Kondo, T. Aoki and I. Hirono

5 : Involvement of WSSV-shrimp homologs in WSSV infectivity in kuruma shrimp: Marsupenaeus japonicus. Antiviral Res., 88, Durand, S. V. and D. V. Lightner2002: Quantitative real time PCR for the measurement of white spot syndrome virus in shrimp. J. Fish Dis., 25, Flegel, T. W.2012: Historic emergence, impact and current status of shrimp pathogens in Asia. J. Invertebr. Pathol., 110, Kono, T., R. Savan, M. Sakai and T. Itami2004: Detection of white spot syndrome virus in shrimp by loop-mediated isothermal amplification. J. Virol. Methods, 115, Kono, T., K. Sonoda, Y. Kitao, T. Mekata, T. Itami and M. Sakai 2009: The expression analysis of innate immune-related genes in kuruma shrimp Penaeus japonicus after DNA vaccination against penaeid rod-shaped DNA virus. Fish Pathol., 44, Kono, T., J. Fall, H. Korenaga, R. Sudhakaran, G. Biswas, T. Mekata, T. Itami and M. Sakai2014: Recombinant VP28 produced by cell-free technique confers protection in Kuruma shrimpmarsupenaeus japonicusagainst white spot syndrome virus. Turkish J. Fish. Aquat. Sci., 14, Leu, J. H., J. M. Tsai, H. C. Wang, A. H. J. Wang, C. H. Wang, G. H. Kou and C. F. Lo2005: The unique stacked rings in the nucleocapsid of the white spot syndrome virus virion are formed by the major structural protein VP664, the largest viral structural protein ever found. J. Virol., 79, Lo, C. F., J. H. Leu, C. H. Ho, C. H. Chen, S. E. Peng, Y. T. Chen, C. M. Chou, P. Y. Yeh, C. J. Huang, H. Y. Chou, C. H. Wang and G. H. Kou1996: Detection of baculovirus associated with white spot syndromewsbvin penaeid shrimps using polymerase chain reaction. Dis. Aquat. Organ., 25, Lo, C. F., S. E. Peng, Y. S. Chang and G. H. Kou2005: White spot syndrome-what we have learned about the virus and the disease. Diseases in Asian aquaculture V. Proceedings of the 5th Symposium on Diseases in Asian Aquaculture. Fish Health Section, Maeda, M., H. Saitoh, E. Mizuki, T. Itami and M. Ohba2004: Replication of white spot syndrome virus in ovarian primary cultures from the kuruma shrimp, Marsupenaeus japonicus. J. Virol. Methods, 116, Mayo, M. A.2002: A summary of taxonomic changes recently approved by ICTV. Arch. Virol., 147, Momoyama, K. and K. Muroga2005: Diseases of cultured kuruma shrimp in Japan: A review. Fish Pathol., 40, Muroga, K.2001: Viral and bacterial diseases of marine fish and shellfish in Japanese hatcheries. Aquaculture, 202, Mushiake, K., M. Arimoto, J. Satoh and K.-i. Mori1998: Detection of PRDV from Wild Adult Kuruma Prawn. Fish Pathol., 33, Namikoshi, A., J. L. Wu, T. Yamashita, T. Nishizawa, T. Nishioka, M. Arimoto and K. Muroga2004: Vaccination trials with Penaeus japonicus to induce resistance to white spot syndrome virus. Aquaculture, 229, , Satoh, J.2012: Studies on prevention measure of white spot disease of kuruma shrimp Marsupenaeus japonicus. Bull Fish Res Agency, Satoh, J., K.-i. Mori, T. Nishioka, K. Hattori, M. Oka and K.-i. Watanabe2006: Disinfection Method for Fertilized Eggs of Kuruma Prawn Penaeus japonicus Using Povidoneiodine. Fish Pathol., 41, PAV 5, PAV30, Thitamadee, S., A. Prachumwat, J. Srisala, P. Jaroenlak, P. V. Salachan, K. Sritunyalucksana, T. W. Flegel and O. Itsathitphaisarn2016: Review of current disease threats for cultivated penaeid shrimp in Asia. Aquaculture, 452, Tsai, J. M., H. C. Wang, J. H. Leu, A. H. J. Wang, Y. Zhuang, P. J. Walker, G. H. Kou and C. F. Lo2006: Identification of the nucleocapsid, tegument, and envelope proteins of the shrimp white spot syndrome virus virion. J. Virol., 80, Utari, H. B., C. Soowannayan, T. W. Flegel, B. Whityachumnarnkul and M. Kruatrachue2017: Variable RNA expression from recently acquired, endogenous viral elements EVEof white spot syndrome viruswssvin shrimp. Dev. Comp. Immunol., 76, Van Hulten, M. C. W., J. Witteveldt, M. Snippe and J. M. Vlak 2001: White spot syndrome virus envelope protein VP28 is involved in the systemic infection of shrimp. Virology, 285, Vlak, J. M., T. W. Flegel, G. H. Kou, D. V. Lightner, C. F. Lo, P. C. Loh and P. J. Walker2005: Family Nimaviridae in Virus taxonomy: classification and nomenclature of viruses: eighth report of the International Committee on the taxonomy of viruses, Witteveldt, J., J. M. Vlak and M. C. W. Van Hulten2004: Protection of Penaeus monodon against white spot syndrome virus using a WSSV subunit vaccine. Fish Shellfish Immunol., 16, Xie, X., L. Xu and F. Yang2006: Proteomic analysis of the major envelope and nucleocapsid proteins of white spot syndrome virus. J. Virol., 80, Yang, F., J. He, X. Lin, Q. Li, D. Pan, X. Zhang and X. Xu2001: Complete genome sequence of the shrimp white spot bacilliform virus. J. Virol., 75,

日本消化器外科学会雑誌第29巻第9号

日本消化器外科学会雑誌第29巻第9号 Table 1 Oligonucleotide primers used for RT-PCR and internal probes used for Southern blot hybridization Cytokine Primer Sequence (5'-3') 5' 3' Internal probe s' 3' Internal probe 5' 3' Internal probe