SrasV1, PascalFrancis, Checkpoint, bibRecord, 000213

Efficient induction of cytotoxic T lymphocytes specific for severe acute respiratory syndrome (SARS)-associated coronavirus by immunization with surface-linked liposomal peptides derived from a non-structural polyprotein 1a

Identifieur interne : 000213 ( PascalFrancis/Checkpoint ); précédent : 000212; suivant : 000214

Efficient induction of cytotoxic T lymphocytes specific for severe acute respiratory syndrome (SARS)-associated coronavirus by immunization with surface-linked liposomal peptides derived from a non-structural polyprotein 1a

Auteurs : Shunsuke Kohyama [Japon] ; Satoshi Ohno [Japon] ; Tatsuya Suda [Japon] ; Maiko Taneichi [Japon] ; Shoichi Yokoyama [Japon] ; Masahito Mori [Japon] ; Akiharu Kobavashi [Japon] ; Hidenori Havashi [Japon] ; Tetsuva Uchida [Japon] ; Masanori Matsui [Japon]

Source :

Antiviral research [ 0166-3542 ] ; 2009.

RBID : Pascal:09-0466845

Descripteurs français

Pascal (Inist)
- Induction, Lymphocyte T cytotoxique, Syndrome respiratoire aigu sévère, Virus syndrome respiratoire aigu sévère, Immunisation, Vaccin, Vaccination, Liposome, Polyprotéine, Prévention, Immunoprophylaxie, Immunité cellulaire, Composé conjugué, Protéine non structurale.
Wicri :
- topic : Vaccin, Vaccination.

English descriptors

KwdEn :
- Cellular immunity, Conjugated compound, Cytotoxic T lymphocyte, Immunization, Immunoprophylaxis, Induction, Liposome, Polyprotein, Prevention, Severe acute respiratory syndrome, Severe acute respiratory syndrome virus, Vaccination, Vaccine.

Abstract

Spike and nucleocapsid are structural proteins of severe acute respiratory syndrome (SARS)-associated coronavirus (SARS-CoV) and major targets for cytotoxic T lymphocytes (CTLs). In contrast, non-structural proteins encoded by two-thirds of viral genome are poorly characterized for cell-mediated immunity. We previously demonstrated that nucleocapsid-derived peptides chemically coupled to the surface of liposomes effectively elicited SARS-CoV-specific CTLs in mice. Here, we attempted to identify HLA-A^*0201-restricted CTL epitopes derived from a non-structural polyprotein 1a (ppla) of SARS-CoV, and investigated whether liposomal peptides derived from pp1a were effective for CTL induction. Out of 30 peptides predicted on computational algorithms, nine peptides could significantly induce interferon gamma (IFN-γ)-producing CD8⁺ T cells in mice. These peptides were coupled to the surface of liposomes, and inoculated into mice. Six liposomal peptides effectively induced IFN-γ-producing CD8⁺ T cells and seven liposomal peptides including the six peptides primed CTLs showing in vivo killing activities. Further, CTLs induced by the seven liposomal peptides lysed an HLA-A^*0201 positive cell line expressing naturally processed, ppla-derived peptides. Of note, one of the liposomal peptides induced high numbers of long-lasting memory CTLs. These data suggest that surface-linked liposomal peptides derived from ppla might offer an efficient CTL-based vaccine against SARS.

Affiliations:

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Le document en format XML

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<author><name sortKey="Kobavashi, Akiharu" sort="Kobavashi, Akiharu" uniqKey="Kobavashi A" first="Akiharu" last="Kobavashi">Akiharu Kobavashi</name>
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<author><name sortKey="Uchida, Tetsuva" sort="Uchida, Tetsuva" uniqKey="Uchida T" first="Tetsuva" last="Uchida">Tetsuva Uchida</name>
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<sourceDesc><biblStruct><analytic><title xml:lang="en" level="a">Efficient induction of cytotoxic T lymphocytes specific for severe acute respiratory syndrome (SARS)-associated coronavirus by immunization with surface-linked liposomal peptides derived from a non-structural polyprotein 1a</title>
<author><name sortKey="Kohyama, Shunsuke" sort="Kohyama, Shunsuke" uniqKey="Kohyama S" first="Shunsuke" last="Kohyama">Shunsuke Kohyama</name>
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<author><name sortKey="Yokoyama, Shoichi" sort="Yokoyama, Shoichi" uniqKey="Yokoyama S" first="Shoichi" last="Yokoyama">Shoichi Yokoyama</name>
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<author><name sortKey="Mori, Masahito" sort="Mori, Masahito" uniqKey="Mori M" first="Masahito" last="Mori">Masahito Mori</name>
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<author><name sortKey="Kobavashi, Akiharu" sort="Kobavashi, Akiharu" uniqKey="Kobavashi A" first="Akiharu" last="Kobavashi">Akiharu Kobavashi</name>
<affiliation wicri:level="1"><inist:fA14 i1="04"><s1>Drug Delivery System Development Division, Nippon Oil and Fat Corporation</s1>
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<author><name sortKey="Havashi, Hidenori" sort="Havashi, Hidenori" uniqKey="Havashi H" first="Hidenori" last="Havashi">Hidenori Havashi</name>
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<author><name sortKey="Uchida, Tetsuva" sort="Uchida, Tetsuva" uniqKey="Uchida T" first="Tetsuva" last="Uchida">Tetsuva Uchida</name>
<affiliation wicri:level="1"><inist:fA14 i1="03"><s1>Department of Safety Research on Blood and Biological Products, National Institute of Infectious Diseases</s1>
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<author><name sortKey="Matsui, Masanori" sort="Matsui, Masanori" uniqKey="Matsui M" first="Masanori" last="Matsui">Masanori Matsui</name>
<affiliation wicri:level="1"><inist:fA14 i1="01"><s1>Department of Microbiology, Faculty of Medicine, Saitama Medical University, Moroyama-cho</s1>
<s2>Iruma-gun, Saitama 350-0495</s2>
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<series><title level="j" type="main">Antiviral research</title>
<title level="j" type="abbreviated">Antivir. res.</title>
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<profileDesc><textClass><keywords scheme="KwdEn" xml:lang="en"><term>Cellular immunity</term>
<term>Conjugated compound</term>
<term>Cytotoxic T lymphocyte</term>
<term>Immunization</term>
<term>Immunoprophylaxis</term>
<term>Induction</term>
<term>Liposome</term>
<term>Polyprotein</term>
<term>Prevention</term>
<term>Severe acute respiratory syndrome</term>
<term>Severe acute respiratory syndrome virus</term>
<term>Vaccination</term>
<term>Vaccine</term>
</keywords>
<keywords scheme="Pascal" xml:lang="fr"><term>Induction</term>
<term>Lymphocyte T cytotoxique</term>
<term>Syndrome respiratoire aigu sévère</term>
<term>Virus syndrome respiratoire aigu sévère</term>
<term>Immunisation</term>
<term>Vaccin</term>
<term>Vaccination</term>
<term>Liposome</term>
<term>Polyprotéine</term>
<term>Prévention</term>
<term>Immunoprophylaxie</term>
<term>Immunité cellulaire</term>
<term>Composé conjugué</term>
<term>Protéine non structurale</term>
</keywords>
<keywords scheme="Wicri" type="topic" xml:lang="fr"><term>Vaccin</term>
<term>Vaccination</term>
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<front><div type="abstract" xml:lang="en">Spike and nucleocapsid are structural proteins of severe acute respiratory syndrome (SARS)-associated coronavirus (SARS-CoV) and major targets for cytotoxic T lymphocytes (CTLs). In contrast, non-structural proteins encoded by two-thirds of viral genome are poorly characterized for cell-mediated immunity. We previously demonstrated that nucleocapsid-derived peptides chemically coupled to the surface of liposomes effectively elicited SARS-CoV-specific CTLs in mice. Here, we attempted to identify HLA-A<sup>*</sup>
0201-restricted CTL epitopes derived from a non-structural polyprotein 1a (ppla) of SARS-CoV, and investigated whether liposomal peptides derived from pp1a were effective for CTL induction. Out of 30 peptides predicted on computational algorithms, nine peptides could significantly induce interferon gamma (IFN-γ)-producing CD8<sup>+</sup>
 T cells in mice. These peptides were coupled to the surface of liposomes, and inoculated into mice. Six liposomal peptides effectively induced IFN-γ-producing CD8<sup>+</sup>
 T cells and seven liposomal peptides including the six peptides primed CTLs showing in vivo killing activities. Further, CTLs induced by the seven liposomal peptides lysed an HLA-A<sup>*</sup>
0201 positive cell line expressing naturally processed, ppla-derived peptides. Of note, one of the liposomal peptides induced high numbers of long-lasting memory CTLs. These data suggest that surface-linked liposomal peptides derived from ppla might offer an efficient CTL-based vaccine against SARS.</div>
</front>
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<fA11 i1="01" i2="1"><s1>KOHYAMA (Shunsuke)</s1>
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<fA11 i1="02" i2="1"><s1>OHNO (Satoshi)</s1>
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<fA11 i1="03" i2="1"><s1>SUDA (Tatsuya)</s1>
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<fA11 i1="05" i2="1"><s1>YOKOYAMA (Shoichi)</s1>
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<fA11 i1="06" i2="1"><s1>MORI (Masahito)</s1>
</fA11>
<fA11 i1="07" i2="1"><s1>KOBAVASHI (Akiharu)</s1>
</fA11>
<fA11 i1="08" i2="1"><s1>HAVASHI (Hidenori)</s1>
</fA11>
<fA11 i1="09" i2="1"><s1>UCHIDA (Tetsuva)</s1>
</fA11>
<fA11 i1="10" i2="1"><s1>MATSUI (Masanori)</s1>
</fA11>
<fA14 i1="01"><s1>Department of Microbiology, Faculty of Medicine, Saitama Medical University, Moroyama-cho</s1>
<s2>Iruma-gun, Saitama 350-0495</s2>
<s3>JPN</s3>
<sZ>1 aut.</sZ>
<sZ>2 aut.</sZ>
<sZ>3 aut.</sZ>
<sZ>10 aut.</sZ>
</fA14>
<fA14 i1="02"><s1>Department of Pathological Biochemistry, Faculty of Pharmaceutical Sciences, Josai University</s1>
<s2>Sakado-city, Saitama 350-0295</s2>
<s3>JPN</s3>
<sZ>1 aut.</sZ>
<sZ>2 aut.</sZ>
<sZ>8 aut.</sZ>
</fA14>
<fA14 i1="03"><s1>Department of Safety Research on Blood and Biological Products, National Institute of Infectious Diseases</s1>
<s2>Musashimurayama-city, Tokyo 208-0011</s2>
<s3>JPN</s3>
<sZ>4 aut.</sZ>
<sZ>9 aut.</sZ>
</fA14>
<fA14 i1="04"><s1>Drug Delivery System Development Division, Nippon Oil and Fat Corporation</s1>
<s2>Tokyo 150-6019</s2>
<s3>JPN</s3>
<sZ>5 aut.</sZ>
<sZ>6 aut.</sZ>
<sZ>7 aut.</sZ>
</fA14>
<fA20><s1>168-177</s1>
</fA20>
<fA21><s1>2009</s1>
</fA21>
<fA23 i1="01"><s0>ENG</s0>
</fA23>
<fA43 i1="01"><s1>INIST</s1>
<s2>18839</s2>
<s5>354000170327240070</s5>
</fA43>
<fA44><s0>0000</s0>
<s1>© 2009 INIST-CNRS. All rights reserved.</s1>
</fA44>
<fA45><s0>1 p.</s0>
</fA45>
<fA47 i1="01" i2="1"><s0>09-0466845</s0>
</fA47>
<fA60><s1>P</s1>
</fA60>
<fA61><s0>A</s0>
</fA61>
<fA64 i1="01" i2="1"><s0>Antiviral research</s0>
</fA64>
<fA66 i1="01"><s0>NLD</s0>
</fA66>
<fC01 i1="01" l="ENG"><s0>Spike and nucleocapsid are structural proteins of severe acute respiratory syndrome (SARS)-associated coronavirus (SARS-CoV) and major targets for cytotoxic T lymphocytes (CTLs). In contrast, non-structural proteins encoded by two-thirds of viral genome are poorly characterized for cell-mediated immunity. We previously demonstrated that nucleocapsid-derived peptides chemically coupled to the surface of liposomes effectively elicited SARS-CoV-specific CTLs in mice. Here, we attempted to identify HLA-A<sup>*</sup>
0201-restricted CTL epitopes derived from a non-structural polyprotein 1a (ppla) of SARS-CoV, and investigated whether liposomal peptides derived from pp1a were effective for CTL induction. Out of 30 peptides predicted on computational algorithms, nine peptides could significantly induce interferon gamma (IFN-γ)-producing CD8<sup>+</sup>
 T cells in mice. These peptides were coupled to the surface of liposomes, and inoculated into mice. Six liposomal peptides effectively induced IFN-γ-producing CD8<sup>+</sup>
 T cells and seven liposomal peptides including the six peptides primed CTLs showing in vivo killing activities. Further, CTLs induced by the seven liposomal peptides lysed an HLA-A<sup>*</sup>
0201 positive cell line expressing naturally processed, ppla-derived peptides. Of note, one of the liposomal peptides induced high numbers of long-lasting memory CTLs. These data suggest that surface-linked liposomal peptides derived from ppla might offer an efficient CTL-based vaccine against SARS.</s0>
</fC01>
<fC02 i1="01" i2="X"><s0>002B02S05</s0>
</fC02>
<fC02 i1="02" i2="X"><s0>002B05C02C</s0>
</fC02>
<fC03 i1="01" i2="X" l="FRE"><s0>Induction</s0>
<s5>01</s5>
</fC03>
<fC03 i1="01" i2="X" l="ENG"><s0>Induction</s0>
<s5>01</s5>
</fC03>
<fC03 i1="01" i2="X" l="SPA"><s0>Inducción</s0>
<s5>01</s5>
</fC03>
<fC03 i1="02" i2="X" l="FRE"><s0>Lymphocyte T cytotoxique</s0>
<s5>02</s5>
</fC03>
<fC03 i1="02" i2="X" l="ENG"><s0>Cytotoxic T lymphocyte</s0>
<s5>02</s5>
</fC03>
<fC03 i1="02" i2="X" l="SPA"><s0>Linfocito T citotóxico</s0>
<s5>02</s5>
</fC03>
<fC03 i1="03" i2="X" l="FRE"><s0>Syndrome respiratoire aigu sévère</s0>
<s2>NM</s2>
<s5>03</s5>
</fC03>
<fC03 i1="03" i2="X" l="ENG"><s0>Severe acute respiratory syndrome</s0>
<s2>NM</s2>
<s5>03</s5>
</fC03>
<fC03 i1="03" i2="X" l="SPA"><s0>Síndrome respiratorio agudo severo</s0>
<s2>NM</s2>
<s5>03</s5>
</fC03>
<fC03 i1="04" i2="X" l="FRE"><s0>Virus syndrome respiratoire aigu sévère</s0>
<s2>NW</s2>
<s5>04</s5>
</fC03>
<fC03 i1="04" i2="X" l="ENG"><s0>Severe acute respiratory syndrome virus</s0>
<s2>NW</s2>
<s5>04</s5>
</fC03>
<fC03 i1="04" i2="X" l="SPA"><s0>Severe acute respiratory syndrome virus</s0>
<s2>NW</s2>
<s5>04</s5>
</fC03>
<fC03 i1="05" i2="X" l="FRE"><s0>Immunisation</s0>
<s5>05</s5>
</fC03>
<fC03 i1="05" i2="X" l="ENG"><s0>Immunization</s0>
<s5>05</s5>
</fC03>
<fC03 i1="05" i2="X" l="SPA"><s0>Inmunización</s0>
<s5>05</s5>
</fC03>
<fC03 i1="06" i2="X" l="FRE"><s0>Vaccin</s0>
<s5>06</s5>
</fC03>
<fC03 i1="06" i2="X" l="ENG"><s0>Vaccine</s0>
<s5>06</s5>
</fC03>
<fC03 i1="06" i2="X" l="SPA"><s0>Vacuna</s0>
<s5>06</s5>
</fC03>
<fC03 i1="07" i2="X" l="FRE"><s0>Vaccination</s0>
<s5>07</s5>
</fC03>
<fC03 i1="07" i2="X" l="ENG"><s0>Vaccination</s0>
<s5>07</s5>
</fC03>
<fC03 i1="07" i2="X" l="SPA"><s0>Vacunación</s0>
<s5>07</s5>
</fC03>
<fC03 i1="08" i2="X" l="FRE"><s0>Liposome</s0>
<s5>09</s5>
</fC03>
<fC03 i1="08" i2="X" l="ENG"><s0>Liposome</s0>
<s5>09</s5>
</fC03>
<fC03 i1="08" i2="X" l="SPA"><s0>Liposoma</s0>
<s5>09</s5>
</fC03>
<fC03 i1="09" i2="X" l="FRE"><s0>Polyprotéine</s0>
<s5>11</s5>
</fC03>
<fC03 i1="09" i2="X" l="ENG"><s0>Polyprotein</s0>
<s5>11</s5>
</fC03>
<fC03 i1="09" i2="X" l="SPA"><s0>Polyproteina</s0>
<s5>11</s5>
</fC03>
<fC03 i1="10" i2="X" l="FRE"><s0>Prévention</s0>
<s5>12</s5>
</fC03>
<fC03 i1="10" i2="X" l="ENG"><s0>Prevention</s0>
<s5>12</s5>
</fC03>
<fC03 i1="10" i2="X" l="SPA"><s0>Prevención</s0>
<s5>12</s5>
</fC03>
<fC03 i1="11" i2="X" l="FRE"><s0>Immunoprophylaxie</s0>
<s5>13</s5>
</fC03>
<fC03 i1="11" i2="X" l="ENG"><s0>Immunoprophylaxis</s0>
<s5>13</s5>
</fC03>
<fC03 i1="11" i2="X" l="SPA"><s0>Inmunoprofilaxia</s0>
<s5>13</s5>
</fC03>
<fC03 i1="12" i2="X" l="FRE"><s0>Immunité cellulaire</s0>
<s5>14</s5>
</fC03>
<fC03 i1="12" i2="X" l="ENG"><s0>Cellular immunity</s0>
<s5>14</s5>
</fC03>
<fC03 i1="12" i2="X" l="SPA"><s0>Inmunidad celular</s0>
<s5>14</s5>
</fC03>
<fC03 i1="13" i2="X" l="FRE"><s0>Composé conjugué</s0>
<s5>15</s5>
</fC03>
<fC03 i1="13" i2="X" l="ENG"><s0>Conjugated compound</s0>
<s5>15</s5>
</fC03>
<fC03 i1="13" i2="X" l="SPA"><s0>Compuesto conjugado</s0>
<s5>15</s5>
</fC03>
<fC03 i1="14" i2="X" l="FRE"><s0>Protéine non structurale</s0>
<s4>INC</s4>
<s5>86</s5>
</fC03>
<fC07 i1="01" i2="X" l="FRE"><s0>Virose</s0>
</fC07>
<fC07 i1="01" i2="X" l="ENG"><s0>Viral disease</s0>
</fC07>
<fC07 i1="01" i2="X" l="SPA"><s0>Virosis</s0>
</fC07>
<fC07 i1="02" i2="X" l="FRE"><s0>Infection</s0>
</fC07>
<fC07 i1="02" i2="X" l="ENG"><s0>Infection</s0>
</fC07>
<fC07 i1="02" i2="X" l="SPA"><s0>Infección</s0>
</fC07>
<fC07 i1="03" i2="X" l="FRE"><s0>Coronavirus</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="03" i2="X" l="ENG"><s0>Coronavirus</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="03" i2="X" l="SPA"><s0>Coronavirus</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="04" i2="X" l="FRE"><s0>Coronaviridae</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="04" i2="X" l="ENG"><s0>Coronaviridae</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="04" i2="X" l="SPA"><s0>Coronaviridae</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="05" i2="X" l="FRE"><s0>Nidovirales</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="05" i2="X" l="ENG"><s0>Nidovirales</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="05" i2="X" l="SPA"><s0>Nidovirales</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="06" i2="X" l="FRE"><s0>Virus</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="06" i2="X" l="ENG"><s0>Virus</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="06" i2="X" l="SPA"><s0>Virus</s0>
<s2>NW</s2>
</fC07>
<fC07 i1="07" i2="X" l="FRE"><s0>Pathologie de l'appareil respiratoire</s0>
<s5>37</s5>
</fC07>
<fC07 i1="07" i2="X" l="ENG"><s0>Respiratory disease</s0>
<s5>37</s5>
</fC07>
<fC07 i1="07" i2="X" l="SPA"><s0>Aparato respiratorio patología</s0>
<s5>37</s5>
</fC07>
<fC07 i1="08" i2="X" l="FRE"><s0>Pathologie des poumons</s0>
<s5>38</s5>
</fC07>
<fC07 i1="08" i2="X" l="ENG"><s0>Lung disease</s0>
<s5>38</s5>
</fC07>
<fC07 i1="08" i2="X" l="SPA"><s0>Pulmón patología</s0>
<s5>38</s5>
</fC07>
<fN21><s1>341</s1>
</fN21>
</pA>
</standard>
</inist>
<affiliations><list><country><li>Japon</li>
</country>
<region><li>Région de Kantō</li>
</region>
<settlement><li>Tokyo</li>
</settlement>
</list>
<tree><country name="Japon"><noRegion><name sortKey="Kohyama, Shunsuke" sort="Kohyama, Shunsuke" uniqKey="Kohyama S" first="Shunsuke" last="Kohyama">Shunsuke Kohyama</name>
</noRegion>
<name sortKey="Havashi, Hidenori" sort="Havashi, Hidenori" uniqKey="Havashi H" first="Hidenori" last="Havashi">Hidenori Havashi</name>
<name sortKey="Kobavashi, Akiharu" sort="Kobavashi, Akiharu" uniqKey="Kobavashi A" first="Akiharu" last="Kobavashi">Akiharu Kobavashi</name>
<name sortKey="Kohyama, Shunsuke" sort="Kohyama, Shunsuke" uniqKey="Kohyama S" first="Shunsuke" last="Kohyama">Shunsuke Kohyama</name>
<name sortKey="Matsui, Masanori" sort="Matsui, Masanori" uniqKey="Matsui M" first="Masanori" last="Matsui">Masanori Matsui</name>
<name sortKey="Mori, Masahito" sort="Mori, Masahito" uniqKey="Mori M" first="Masahito" last="Mori">Masahito Mori</name>
<name sortKey="Ohno, Satoshi" sort="Ohno, Satoshi" uniqKey="Ohno S" first="Satoshi" last="Ohno">Satoshi Ohno</name>
<name sortKey="Ohno, Satoshi" sort="Ohno, Satoshi" uniqKey="Ohno S" first="Satoshi" last="Ohno">Satoshi Ohno</name>
<name sortKey="Suda, Tatsuya" sort="Suda, Tatsuya" uniqKey="Suda T" first="Tatsuya" last="Suda">Tatsuya Suda</name>
<name sortKey="Taneichi, Maiko" sort="Taneichi, Maiko" uniqKey="Taneichi M" first="Maiko" last="Taneichi">Maiko Taneichi</name>
<name sortKey="Uchida, Tetsuva" sort="Uchida, Tetsuva" uniqKey="Uchida T" first="Tetsuva" last="Uchida">Tetsuva Uchida</name>
<name sortKey="Yokoyama, Shoichi" sort="Yokoyama, Shoichi" uniqKey="Yokoyama S" first="Shoichi" last="Yokoyama">Shoichi Yokoyama</name>
</country>
</tree>
</affiliations>
</record>

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   |wiki=    Sante
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   |texte=   Efficient induction of cytotoxic T lymphocytes specific for severe acute respiratory syndrome (SARS)-associated coronavirus by immunization with surface-linked liposomal peptides derived from a non-structural polyprotein 1a
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Efficient induction of cytotoxic T lymphocytes specific for severe acute respiratory syndrome (SARS)-associated coronavirus by immunization with surface-linked liposomal peptides derived from a non-structural polyprotein 1a

Efficient induction of cytotoxic T lymphocytes specific for severe acute respiratory syndrome (SARS)-associated coronavirus by immunization with surface-linked liposomal peptides derived from a non-structural polyprotein 1a

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