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A New Nucleopolyhedrovirus Strain (Ldmnpv-Like Virus) with a Defective Fp25 Gene from Lymantria Xylina (Lepidoptera: Lymantriidae) in Taiwan

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A New Nucleopolyhedrovirus Strain (Ldmnpv-Like Virus) with a Defective Fp25 Gene from Lymantria Xylina (Lepidoptera: Lymantriidae) in Taiwan Journal of Invertebrate Pathology 102 (2009) 110–119 Contents lists available at ScienceDirect Journal of Invertebrate Pathology journal homepage: www.elsevier.com/locate/yjipa A new nucleopolyhedrovirus strain (LdMNPV-like virus) with a defective fp25 gene from Lymantria xylina (Lepidoptera: Lymantriidae) in Taiwan Yu-Shin Nai a,1, Tai-Chuan Wang a,1, Yun-Ru Chen a,1, Chu-Fang Lo b,*, Chung-Hsiung Wang a,b,* a Department of Entomology, National Taiwan University, Taipei, Taiwan, ROC b Department of Zoology, National Taiwan University, Taipei, Taiwan, ROC article info abstract Article history: A new multiple nucleopolyhedrovirus strain was isolated from casuarina moth, Lymantria xylina Swinhoe, Received 1 August 2008 (Lepidoptera: Lymantriidae) in Taiwan. This Lymantria-derived virus can be propagated in IPLB-LD-652Y Accepted 13 July 2009 and NTU-LY cell lines and showed a few polyhedra (occlusion bodies) CPE in the infected cells. The Available online 17 July 2009 restriction fragment length polymorphism (RFLP) profiles of whole genome indicated that this virus is distinct from LyxyMNPV and the virus genome size was approximately 139 kbps, which was smaller than Keywords: that of LyxyMNPV. The molecular phylogenetic analyses of three important genes (polyhedrin, lef-8 and LyxyMNPV lef-9) were performed. Polyhedrin, LEF-8 and LEF-9 putative amino acid analyses of this virus revealed LdMNPV-like virus that this virus belongs to Group II NPV and closely related to LdMNPV than to LyxyMNPV. The phyloge- LdMNPV Fp25k mutant netic distance analysis was further clarified the relationship to LdMNPV and this virus provisionally RFLP named LdMNPV-like virus. A significant deletion of a 44 bp sequence found in LdMNPV-like virus was noted in the fp25k sequences of LdMNPV and LyxyMNPV and may play an important role in the few poly- hedra CPE. In ultrastructural observations, the nuclei of the infected LD host cells contained large occlu- sion bodies (OBs), and few OBs, which presented as one or two OBs in a nucleus that was otherwise filled with free nuclocapsids and virions. We concluded that this LdMNPV-like virus is a new LdMNPV strain from L. xylina. Crown Copyright Ó 2009 Published by Elsevier Inc. All rights reserved. 1. Introduction chemical insecticides are still used to control L. xylina caterpil- lars, but this method causes serious environmental impacts, as The casuarina moth, Lymantria xylina Swinehoe (Lepidoptera: well as raising human health concerns. Lymantriidae), is a herbivore that feeds on casuarina (Casuarina A nucleopolyhedrosis epizootic of L. xylina larvae occurs each equisetifolia), guava (Psidium guajava L.), lingan (Euphoria longana year from spring to early summer in Taiwan and mainland China, Lam.), lichi (lychee, Litchi chinensis Sonn.), acacia (Acacia confusa) and the key pathogen was found to be L. xylina multiple nucleo- forests and more than 60 other species of host plants (Chao polyhedrovirus (LyxyMNPV) (Cheng et al., 1987; Liang et al., et al., 1996; Shen et al., 2003; Shen et al., 2006). It is native to 1986; Wu and Wang, 2005; Xiao, 1992; Yu et al., 1997). Lyx- Taiwan, Japan, India, and the eastern coast of mainland China yMNPV’s high virulence to L. xylina suggested that it would be a (Chao et al., 1996; Matsumura, 1933; Xiao, 1992), but in Taiwan, promising agent to include in integrated pest management (IPM) in the last 30 years, as forest was completed to crops, the moth programs for the biological control of L. xylina. According to Wu expanded its range into these newly established agricultural and Wang (2005), LyxyMNPV is closely related to LdMNPV and be- areas and simultaneously expanded its host plant range (Chao longs to group II NPV. et al., 1996). Since 2001, infestations of this moth have damage In the present paper, we characterize a new Lymantria-de- to the Taiwan fruit industry, and L. xylina is now considered to rived virus that was first isolated from infected larva of L. xylina be a significant pest that is subject to quarantine in Taiwan (Yu et al., 1997). This new strain showed a high identity to the (Chao, 2002). L. xylina is phylogenetically closely related to the polyhedrin sequence of LdMNPV, and the in vitro infectivity of gypsy moth, Lymantria dispar, a very important forest pest in this strain in LY and LD cells was similar to that of wild-type Europe and North America (Solter and Hajek, 2009). In Taiwan, (WT) LyxyMNPV (Wu and Wang, 2005, 2006). Over 800 different baculovirus isolates from a variety of invertebrates have been re- ported. Genotypic variants or distinct isolates have been isolated * Corresponding authors. Fax: +886 2 27364329 (C.H. Wang). E-mail address: [email protected] (C.-H. Wang). from the same host species from sympatric or allopatric areas 1 Equal contribution to this paper. (Lee and Miller, 1978; Gettig and McCarthy, 1982; Cherry and 0022-2011/$ - see front matter Crown Copyright Ó 2009 Published by Elsevier Inc. All rights reserved. doi:10.1016/j.jip.2009.07.004 Y.-S. Nai et al. / Journal of Invertebrate Pathology 102 (2009) 110–119 111 Summers, 1985; Lee and Lee, 1988; Maeda et al., 1990; Slavicek (CPE) of the FP variants is usually defined as less than 10 OBs et al., 1992; Weitzman et al., 1992), whereas few polyhedra (FP) in the infected cells, FP variants not only show reduced numbers (few occlusion bodies or OBs) and many polyhedra (MP) or OB of OBs in the infected cell, but also OBs with few or no virions variants of NPVs usually occur during the serial cell culture pas- occluded. In contrast, FP variants increase the yield of budded sage at high M.O.I. (multiplicity of infection) rates (Tramper and virus (BV) (Harrison and Summers, 1995; Bull et al., 2003; Wu Vlak, 1986). et al., 2005). Since Hink and Vail (1973) first observed FP mutants in the The FP CPE is mainly the result of a deficient fp25k gene in the AcMNPV-infected Trichopusia ni cells, the common properties of baculoviral genome. The fp25k mutants usually produce a great the FP CPE have been studied in several baculoviruses. In vitro number of budded virus (BV) into the cell culture medium, and serial passage of baculovirus will frequently lead to production virus progeny at 72 h post infection (pi) are still observed budding of FP variants (Fraser et al., 1983). The cyptopathoic effects at the plasma membrane, whereas in wild-type (WT) NPVs, BV pro- Table 1 Baculoviruses Polyhedrin, LEF-8 and LEF-9 amino acid sequences used in this study. Group Virus name Accession number References Polyhedrin LEF-8 LEF-9 Lepidopteran NPV-I AcMNPV NC_001623 NP_054079 NP_054092 Ayres et al. (1994) AgMNPV YP_803395 YP_803445 YP_803456 Oliveira et al. (2006) AnpeNPV YP_610970 YP_611067 YP_611058 Nie et al. (2007) BmNPV NP_047414 NP_047454 NP_047466 Gomi et al. (1999) CfMNPV NP_848313 NP_848359 NP_848370 de Jong et al. (2005) CfDE FNPV NP_932610 NP_932656 NP_932667 Lauzon et al. (2005) Epp oNPV NP_203170 NP_203214 NP_203225 Hyink et al. (2002) HycuNP V YP_473189 YP_473277 YP_473277 Ikeda et at. (2006) MaviNPV YP_950731 YP_950765 YP_950776 Chen et al. (2008) OpMNPV NP_046159 NP_046210 NP_046221 Ahrens et al. (1997) RoMNPV NP_702998 NP_703040 NP_703052 Harrison and Bonning (2003) Lepidopteran NPV-II AdhoNPV NP_818648 NP_818696 NP_818682 Nakai et al. (2003) AgseNPV YP_529671 YP_529791 YP_529775 Jakubowska et al. (2006) ChchNPV YP_249605 YP_249641 YP_249656 van Oers et at. (2005) C1biNPV YP_717539 YP_717570 YP_717585 Wang et al. (unpublished) EcobNPV YP_874194 YP_874225 YP_874239 Ma et al. (2006) HearNP V (G4) NP_075070 NP_075107 NP_075124 Chen et al. (2001) HearNP V (Cl) NP_203559 NP_203594 NP_203610 Zhang et al. (2005) HzSNPV NP_542624 NP_542661 NP_542679 Chen et al. (2002) LdMNPV NP_047637 NP_047687 NP_047701 Kuzio et al. (1999) LsNPV YP_758298 YP_758340 YP_758375 Xiao and Qi (2007) LyxyMNPV AAW28 857 ACL31659 ACL31660 Wu and Wang (2005) and This study LdMNPV-like virus ACL31661 ACL31658 ACL31657 This study MacoNPV (A) NP_613084 NP_613224 NP_613207 Li et al. (2002b) MacoNPV (B) NP_689176 NP_689314 NP_689297 Li et al. (2002a) SeMNPV NP_037761 NP_037872 NP_037857 IJkel et al.(1999) SfMNP V YP_001036294 YP_001036405 YP_001036389 Harrison et al. (2008) Sp1tNPV NP_258269 NP_258306 NP_258327 Pang et al. (2001) TnSNPV YP308889 YP_308923 YP_308939 Willis et al. (2005) Lepidopteran GV CpGV NP_148785 NP_148915 NP_148901 Luque et al. (2001) Hymenopteran NPV Nea bNPV YP_667849 YP_667931 YP_667887 Duffy et al. (2006) Ne1eNPV YP_025198 YP_025279 YP_025237 Lauzon et al. (2004) NeseNPV YP_025108 YP_025188 YP_025147 Garcia-Manniak et al. (2004) Fig. 1. Cytopathic effects (CPE) of IPLB-LD-652Y cells infected with LyxyMNPV and LdMNPV-like virus at 7 days pi showing: (A) MP (multiple polyhedra) and FP (few polyhedra) found in the LyxyMNPV-infected LD cells; (B) FP only in the LdMNPV-like virus-infected LD cells; and (C) uninfected LD cells. Bar: 25 lm. 112 Y.-S. Nai et al. / Journal of Invertebrate Pathology 102 (2009) 110–119 duction decreases to barely detectable levels (Braunagel et al., Lymantria-derived virus system will provide the in vivo and in vitro 1999; Wu et al., 2005). co-infection system to evaluate the pathogenesis of simultaneous In nature, the population of a WT genotype is highly predomi- infection in host cells. nant over that of the occlusion-negative genotype and finally the population of occlusion-negative genotype will be lost (Bull et al., 2003). In laboratory, the occlusion-negative genotype can be main- 2. Materials and methods tained in the susceptible cells. Usually, FP mutant virus has been observed to persist in a stable (Bull et al., 2003). The process of 2.1. Insect cell lines and viruses such co-occlusion with the WT and the occlusion-negative geno- types had been suggested as a strategy for application of the genet- IPLB-LD-652Y cells (L. dispar cell line; Goodwin et al., 1978) ically modification baculovirus for insecticidal purposes (Wood and NTU-LY cells (L.
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