Proanthocyanidin-enriched extract from Myrothamnus flabellifolia Welw. exerts antiviral activity against herpes simplex virus type 1 by inhibition of viral adsorption and penetration
Extracts from the aerial parts of the South African resurrection plant Myrothamnus flabellifolia Welw. have been used traditionally against infections of the upper respiratory tract and skin diseases. A polyphenol-enriched extract was investigated for potential antiviral effects against herpes simpl...
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| Veröffentlicht in: | Journal of ethnopharmacology 2011-03, Vol.134 (2), p.468-474 |
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| creator | Gescher, Kirsten Kühn, Joachim Lorentzen, Eva Hafezi, Wali Derksen, Andrea Deters, Alexandra Hensel, Andreas |
| description | Extracts from the aerial parts of the South African resurrection plant Myrothamnus flabellifolia Welw. have been used traditionally against infections of the upper respiratory tract and skin diseases. A polyphenol-enriched extract was investigated for potential antiviral effects against herpes simplex virus type 1 (HSV-1) and adenovirus, and the underlying mode of action was to be studied.
Antiviral effects of an acetone–water extract (MF) from Myrothamnus flabellifolia on HSV-1 and adenovirus type 3 were tested in infected Vero cells by plaque reduction assay, MTT test and immunofluorescence. The influence of the extract on the HSV-1 envelope glycoprotein D was shown by Western blot. Organotypic full thickness skin models consisting of multilayer skin equivalents were used for the investigation of MF effects on HSV-1 replication.
MF exhibited strong antiviral activity against HSV-1. The HSV-1-specific inhibitory concentration (IC50) was determined as 0.4μg/mL and the cytotoxic concentration (CC50) against Vero cells as 50μg/mL. A selectivity index (SI) (ratio of CC50 to IC50) of approximately 120 was calculated when MF was added to the virus inoculum for 1h at 37°C prior to infection. The replication of adenovirus 3 was not affected by MF.
MF abolished virus entry into the host cell by blocking viral attachment to the cell surface. When added after attachment at a concentration of >6μg/mL, the extract also inhibited penetration of HSV-1 into the host cell. Polyphenolic compounds from MF directly interacted with viral particles, leading to the oligomerisation of envelope proteins as demonstrated for the essential viral glycoprotein D (gD). Using organotypic full thickness tissue cultures, it was shown that treatment of HSV-1 infected cultures with the MF resulted in reduced viral spread.
A polyphenol-enriched extract from Myrothamnus flabellifolia strongly acts against HSV-1 by blocking viral entry into the cells. |
| doi_str_mv | 10.1016/j.jep.2010.12.038 |
| format | Article |
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Antiviral effects of an acetone–water extract (MF) from Myrothamnus flabellifolia on HSV-1 and adenovirus type 3 were tested in infected Vero cells by plaque reduction assay, MTT test and immunofluorescence. The influence of the extract on the HSV-1 envelope glycoprotein D was shown by Western blot. Organotypic full thickness skin models consisting of multilayer skin equivalents were used for the investigation of MF effects on HSV-1 replication.
MF exhibited strong antiviral activity against HSV-1. The HSV-1-specific inhibitory concentration (IC50) was determined as 0.4μg/mL and the cytotoxic concentration (CC50) against Vero cells as 50μg/mL. A selectivity index (SI) (ratio of CC50 to IC50) of approximately 120 was calculated when MF was added to the virus inoculum for 1h at 37°C prior to infection. The replication of adenovirus 3 was not affected by MF.
MF abolished virus entry into the host cell by blocking viral attachment to the cell surface. When added after attachment at a concentration of >6μg/mL, the extract also inhibited penetration of HSV-1 into the host cell. Polyphenolic compounds from MF directly interacted with viral particles, leading to the oligomerisation of envelope proteins as demonstrated for the essential viral glycoprotein D (gD). Using organotypic full thickness tissue cultures, it was shown that treatment of HSV-1 infected cultures with the MF resulted in reduced viral spread.
A polyphenol-enriched extract from Myrothamnus flabellifolia strongly acts against HSV-1 by blocking viral entry into the cells.</description><identifier>ISSN: 0378-8741</identifier><identifier>EISSN: 1872-7573</identifier><identifier>DOI: 10.1016/j.jep.2010.12.038</identifier><identifier>PMID: 21211557</identifier><identifier>CODEN: JOETD7</identifier><language>eng</language><publisher>Shannon: Elsevier Ireland Ltd</publisher><subject>Adenoviridae - drug effects ; Adenoviridae Infections - microbiology ; Adenovirus ; Adsorption ; aerial parts ; Animals ; Antiviral ; Antiviral activity ; Antiviral Agents - pharmacology ; Antiviral Agents - therapeutic use ; antiviral properties ; Attachment ; Biological and medical sciences ; Cell culture ; Cell Line ; Cell surface ; Cercopithecus aethiops ; Cytotoxicity ; Envelope protein ; fluorescent antibody technique ; gD protein ; General pharmacology ; glycoprotein D ; glycoproteins ; Herpes Simplex - drug therapy ; Herpes Simplex - microbiology ; Herpes simplex virus 1 ; Herpes simplex virus type-1 ; Herpesvirus 1, Human - chemistry ; Herpesvirus 1, Human - drug effects ; Herpesvirus 1, Human - pathogenicity ; hosts ; Human herpesvirus 1 ; Humans ; Immunofluorescence ; Infection ; Inhibitory Concentration 50 ; Inoculum ; Keratinocytes - drug effects ; Keratinocytes - microbiology ; Magnoliopsida - chemistry ; mechanism of action ; Medical sciences ; Myrothamnaceae ; Myrothamnus ; Myrothamnus flabellifolia Welw ; Organotypic skin model ; Penetration ; Pharmacognosy. Homeopathy. Health food ; Pharmacology. Drug treatments ; Phytotherapy ; Plant Components, Aerial ; Plant diseases ; Plant Extracts - pharmacology ; Plant Extracts - therapeutic use ; Plaques ; Polyphenols ; Proanthocyanidins - pharmacology ; Proanthocyanidins - therapeutic use ; Replication ; Respiratory tract ; respiratory tract diseases ; Skin - drug effects ; Skin - microbiology ; Skin diseases ; Tissue culture ; Vero Cells ; Viral Envelope Proteins - chemistry ; virion ; Virus Integration - drug effects ; viruses ; Western blotting</subject><ispartof>Journal of ethnopharmacology, 2011-03, Vol.134 (2), p.468-474</ispartof><rights>2011 Elsevier Ireland Ltd</rights><rights>2015 INIST-CNRS</rights><rights>Copyright © 2011 Elsevier Ireland Ltd. All rights reserved.</rights><lds50>peer_reviewed</lds50><woscitedreferencessubscribed>false</woscitedreferencessubscribed><citedby>FETCH-LOGICAL-c438t-bd351896566b15121401c05c4e7087ad831e6ec24d141b3f41a3a76c290793e33</citedby><cites>FETCH-LOGICAL-c438t-bd351896566b15121401c05c4e7087ad831e6ec24d141b3f41a3a76c290793e33</cites></display><links><openurl>$$Topenurl_article</openurl><openurlfulltext>$$Topenurlfull_article</openurlfulltext><thumbnail>$$Tsyndetics_thumb_exl</thumbnail><linktohtml>$$Uhttps://www.sciencedirect.com/science/article/pii/S0378874110009141$$EHTML$$P50$$Gelsevier$$H</linktohtml><link.rule.ids>314,776,780,3537,27901,27902,65306</link.rule.ids><backlink>$$Uhttp://pascal-francis.inist.fr/vibad/index.php?action=getRecordDetail&idt=23976350$$DView record in Pascal Francis$$Hfree_for_read</backlink><backlink>$$Uhttps://www.ncbi.nlm.nih.gov/pubmed/21211557$$D View this record in MEDLINE/PubMed$$Hfree_for_read</backlink></links><search><creatorcontrib>Gescher, Kirsten</creatorcontrib><creatorcontrib>Kühn, Joachim</creatorcontrib><creatorcontrib>Lorentzen, Eva</creatorcontrib><creatorcontrib>Hafezi, Wali</creatorcontrib><creatorcontrib>Derksen, Andrea</creatorcontrib><creatorcontrib>Deters, Alexandra</creatorcontrib><creatorcontrib>Hensel, Andreas</creatorcontrib><title>Proanthocyanidin-enriched extract from Myrothamnus flabellifolia Welw. exerts antiviral activity against herpes simplex virus type 1 by inhibition of viral adsorption and penetration</title><title>Journal of ethnopharmacology</title><addtitle>J Ethnopharmacol</addtitle><description>Extracts from the aerial parts of the South African resurrection plant Myrothamnus flabellifolia Welw. have been used traditionally against infections of the upper respiratory tract and skin diseases. A polyphenol-enriched extract was investigated for potential antiviral effects against herpes simplex virus type 1 (HSV-1) and adenovirus, and the underlying mode of action was to be studied.
Antiviral effects of an acetone–water extract (MF) from Myrothamnus flabellifolia on HSV-1 and adenovirus type 3 were tested in infected Vero cells by plaque reduction assay, MTT test and immunofluorescence. The influence of the extract on the HSV-1 envelope glycoprotein D was shown by Western blot. Organotypic full thickness skin models consisting of multilayer skin equivalents were used for the investigation of MF effects on HSV-1 replication.
MF exhibited strong antiviral activity against HSV-1. The HSV-1-specific inhibitory concentration (IC50) was determined as 0.4μg/mL and the cytotoxic concentration (CC50) against Vero cells as 50μg/mL. A selectivity index (SI) (ratio of CC50 to IC50) of approximately 120 was calculated when MF was added to the virus inoculum for 1h at 37°C prior to infection. The replication of adenovirus 3 was not affected by MF.
MF abolished virus entry into the host cell by blocking viral attachment to the cell surface. When added after attachment at a concentration of >6μg/mL, the extract also inhibited penetration of HSV-1 into the host cell. Polyphenolic compounds from MF directly interacted with viral particles, leading to the oligomerisation of envelope proteins as demonstrated for the essential viral glycoprotein D (gD). Using organotypic full thickness tissue cultures, it was shown that treatment of HSV-1 infected cultures with the MF resulted in reduced viral spread.
A polyphenol-enriched extract from Myrothamnus flabellifolia strongly acts against HSV-1 by blocking viral entry into the cells.</description><subject>Adenoviridae - drug effects</subject><subject>Adenoviridae Infections - microbiology</subject><subject>Adenovirus</subject><subject>Adsorption</subject><subject>aerial parts</subject><subject>Animals</subject><subject>Antiviral</subject><subject>Antiviral activity</subject><subject>Antiviral Agents - pharmacology</subject><subject>Antiviral Agents - therapeutic use</subject><subject>antiviral properties</subject><subject>Attachment</subject><subject>Biological and medical sciences</subject><subject>Cell culture</subject><subject>Cell Line</subject><subject>Cell surface</subject><subject>Cercopithecus aethiops</subject><subject>Cytotoxicity</subject><subject>Envelope protein</subject><subject>fluorescent antibody technique</subject><subject>gD protein</subject><subject>General pharmacology</subject><subject>glycoprotein D</subject><subject>glycoproteins</subject><subject>Herpes Simplex - drug therapy</subject><subject>Herpes Simplex - microbiology</subject><subject>Herpes simplex virus 1</subject><subject>Herpes simplex virus type-1</subject><subject>Herpesvirus 1, Human - chemistry</subject><subject>Herpesvirus 1, Human - drug effects</subject><subject>Herpesvirus 1, Human - pathogenicity</subject><subject>hosts</subject><subject>Human herpesvirus 1</subject><subject>Humans</subject><subject>Immunofluorescence</subject><subject>Infection</subject><subject>Inhibitory Concentration 50</subject><subject>Inoculum</subject><subject>Keratinocytes - drug effects</subject><subject>Keratinocytes - microbiology</subject><subject>Magnoliopsida - chemistry</subject><subject>mechanism of action</subject><subject>Medical sciences</subject><subject>Myrothamnaceae</subject><subject>Myrothamnus</subject><subject>Myrothamnus flabellifolia Welw</subject><subject>Organotypic skin model</subject><subject>Penetration</subject><subject>Pharmacognosy. Homeopathy. Health food</subject><subject>Pharmacology. Drug treatments</subject><subject>Phytotherapy</subject><subject>Plant Components, Aerial</subject><subject>Plant diseases</subject><subject>Plant Extracts - pharmacology</subject><subject>Plant Extracts - therapeutic use</subject><subject>Plaques</subject><subject>Polyphenols</subject><subject>Proanthocyanidins - pharmacology</subject><subject>Proanthocyanidins - therapeutic use</subject><subject>Replication</subject><subject>Respiratory tract</subject><subject>respiratory tract diseases</subject><subject>Skin - drug effects</subject><subject>Skin - microbiology</subject><subject>Skin diseases</subject><subject>Tissue culture</subject><subject>Vero Cells</subject><subject>Viral Envelope Proteins - chemistry</subject><subject>virion</subject><subject>Virus Integration - drug effects</subject><subject>viruses</subject><subject>Western blotting</subject><issn>0378-8741</issn><issn>1872-7573</issn><fulltext>true</fulltext><rsrctype>article</rsrctype><creationdate>2011</creationdate><recordtype>article</recordtype><sourceid>EIF</sourceid><recordid>eNqFkU2O1DAQhSMEYpqBA7ABbxCrNC7bidNihUb8SYNAghFLy3EqE7cSO9juYXIxzodDN7CDlV3W915V-RXFY6BboFC_2G_3OG8ZXWu2pby5U2ygkayUleR3iw3lsikbKeCseBDjnlIqQdD7xRkDBlBVclP8-BS8dmnwZtHOdtaV6II1A3YEb1PQJpE--Il8WIJPg57cIZJ-1C2Oo-39aDX5iuP3bYYxpEiylb2xQY8kK_MtLURfa-tiIgOGGSOJdppHvCWZylZpmZEAaRdi3WBbm6x3xPfk5NFFH-Zfb9p1ZEaHeaS1fljc6_UY8dHpPC-u3rz-cvGuvPz49v3Fq8vSCN6ksu14Bc2uruq6hSovLSgYWhmBkjZSdw0HrNEw0YGAlvcCNNeyNmxH5Y4j5-fF86PvHPy3A8akJhtNXl479IeodpRxwQWw_5JNJYExWq-ecCRN8DEG7NUc7KTDooCqNVe1VzlXteaqgKmca9Y8Obkf2gm7P4rfQWbg2QnQ0eixD9oZG_9yfCdrXtHMPT1yvfZKX4fMXH3OnQSl0ACt1lYvjwTmf72xGFQ0Fp3BzgY0SXXe_mPQn1lkzSw</recordid><startdate>20110324</startdate><enddate>20110324</enddate><creator>Gescher, Kirsten</creator><creator>Kühn, Joachim</creator><creator>Lorentzen, Eva</creator><creator>Hafezi, Wali</creator><creator>Derksen, Andrea</creator><creator>Deters, Alexandra</creator><creator>Hensel, Andreas</creator><general>Elsevier Ireland Ltd</general><general>Elsevier</general><scope>FBQ</scope><scope>IQODW</scope><scope>CGR</scope><scope>CUY</scope><scope>CVF</scope><scope>ECM</scope><scope>EIF</scope><scope>NPM</scope><scope>AAYXX</scope><scope>CITATION</scope><scope>7X8</scope><scope>7T7</scope><scope>7U9</scope><scope>8FD</scope><scope>C1K</scope><scope>FR3</scope><scope>H94</scope><scope>P64</scope></search><sort><creationdate>20110324</creationdate><title>Proanthocyanidin-enriched extract from Myrothamnus flabellifolia Welw. exerts antiviral activity against herpes simplex virus type 1 by inhibition of viral adsorption and penetration</title><author>Gescher, Kirsten ; Kühn, Joachim ; Lorentzen, Eva ; Hafezi, Wali ; Derksen, Andrea ; Deters, Alexandra ; Hensel, Andreas</author></sort><facets><frbrtype>5</frbrtype><frbrgroupid>cdi_FETCH-LOGICAL-c438t-bd351896566b15121401c05c4e7087ad831e6ec24d141b3f41a3a76c290793e33</frbrgroupid><rsrctype>articles</rsrctype><prefilter>articles</prefilter><language>eng</language><creationdate>2011</creationdate><topic>Adenoviridae - drug effects</topic><topic>Adenoviridae Infections - microbiology</topic><topic>Adenovirus</topic><topic>Adsorption</topic><topic>aerial parts</topic><topic>Animals</topic><topic>Antiviral</topic><topic>Antiviral activity</topic><topic>Antiviral Agents - pharmacology</topic><topic>Antiviral Agents - therapeutic use</topic><topic>antiviral properties</topic><topic>Attachment</topic><topic>Biological and medical sciences</topic><topic>Cell culture</topic><topic>Cell Line</topic><topic>Cell surface</topic><topic>Cercopithecus aethiops</topic><topic>Cytotoxicity</topic><topic>Envelope protein</topic><topic>fluorescent antibody technique</topic><topic>gD protein</topic><topic>General pharmacology</topic><topic>glycoprotein D</topic><topic>glycoproteins</topic><topic>Herpes Simplex - drug therapy</topic><topic>Herpes Simplex - microbiology</topic><topic>Herpes simplex virus 1</topic><topic>Herpes simplex virus type-1</topic><topic>Herpesvirus 1, Human - chemistry</topic><topic>Herpesvirus 1, Human - drug effects</topic><topic>Herpesvirus 1, Human - pathogenicity</topic><topic>hosts</topic><topic>Human herpesvirus 1</topic><topic>Humans</topic><topic>Immunofluorescence</topic><topic>Infection</topic><topic>Inhibitory Concentration 50</topic><topic>Inoculum</topic><topic>Keratinocytes - drug effects</topic><topic>Keratinocytes - microbiology</topic><topic>Magnoliopsida - chemistry</topic><topic>mechanism of action</topic><topic>Medical sciences</topic><topic>Myrothamnaceae</topic><topic>Myrothamnus</topic><topic>Myrothamnus flabellifolia Welw</topic><topic>Organotypic skin model</topic><topic>Penetration</topic><topic>Pharmacognosy. Homeopathy. Health food</topic><topic>Pharmacology. Drug treatments</topic><topic>Phytotherapy</topic><topic>Plant Components, Aerial</topic><topic>Plant diseases</topic><topic>Plant Extracts - pharmacology</topic><topic>Plant Extracts - therapeutic use</topic><topic>Plaques</topic><topic>Polyphenols</topic><topic>Proanthocyanidins - pharmacology</topic><topic>Proanthocyanidins - therapeutic use</topic><topic>Replication</topic><topic>Respiratory tract</topic><topic>respiratory tract diseases</topic><topic>Skin - drug effects</topic><topic>Skin - microbiology</topic><topic>Skin diseases</topic><topic>Tissue culture</topic><topic>Vero Cells</topic><topic>Viral Envelope Proteins - chemistry</topic><topic>virion</topic><topic>Virus Integration - drug effects</topic><topic>viruses</topic><topic>Western blotting</topic><toplevel>peer_reviewed</toplevel><toplevel>online_resources</toplevel><creatorcontrib>Gescher, Kirsten</creatorcontrib><creatorcontrib>Kühn, Joachim</creatorcontrib><creatorcontrib>Lorentzen, Eva</creatorcontrib><creatorcontrib>Hafezi, Wali</creatorcontrib><creatorcontrib>Derksen, Andrea</creatorcontrib><creatorcontrib>Deters, Alexandra</creatorcontrib><creatorcontrib>Hensel, Andreas</creatorcontrib><collection>AGRIS</collection><collection>Pascal-Francis</collection><collection>Medline</collection><collection>MEDLINE</collection><collection>MEDLINE (Ovid)</collection><collection>MEDLINE</collection><collection>MEDLINE</collection><collection>PubMed</collection><collection>CrossRef</collection><collection>MEDLINE - Academic</collection><collection>Industrial and Applied Microbiology Abstracts (Microbiology A)</collection><collection>Virology and AIDS Abstracts</collection><collection>Technology Research Database</collection><collection>Environmental Sciences and Pollution Management</collection><collection>Engineering Research Database</collection><collection>AIDS and Cancer Research Abstracts</collection><collection>Biotechnology and BioEngineering Abstracts</collection><jtitle>Journal of ethnopharmacology</jtitle></facets><delivery><delcategory>Remote Search Resource</delcategory><fulltext>fulltext</fulltext></delivery><addata><au>Gescher, Kirsten</au><au>Kühn, Joachim</au><au>Lorentzen, Eva</au><au>Hafezi, Wali</au><au>Derksen, Andrea</au><au>Deters, Alexandra</au><au>Hensel, Andreas</au><format>journal</format><genre>article</genre><ristype>JOUR</ristype><atitle>Proanthocyanidin-enriched extract from Myrothamnus flabellifolia Welw. exerts antiviral activity against herpes simplex virus type 1 by inhibition of viral adsorption and penetration</atitle><jtitle>Journal of ethnopharmacology</jtitle><addtitle>J Ethnopharmacol</addtitle><date>2011-03-24</date><risdate>2011</risdate><volume>134</volume><issue>2</issue><spage>468</spage><epage>474</epage><pages>468-474</pages><issn>0378-8741</issn><eissn>1872-7573</eissn><coden>JOETD7</coden><abstract>Extracts from the aerial parts of the South African resurrection plant Myrothamnus flabellifolia Welw. have been used traditionally against infections of the upper respiratory tract and skin diseases. A polyphenol-enriched extract was investigated for potential antiviral effects against herpes simplex virus type 1 (HSV-1) and adenovirus, and the underlying mode of action was to be studied.
Antiviral effects of an acetone–water extract (MF) from Myrothamnus flabellifolia on HSV-1 and adenovirus type 3 were tested in infected Vero cells by plaque reduction assay, MTT test and immunofluorescence. The influence of the extract on the HSV-1 envelope glycoprotein D was shown by Western blot. Organotypic full thickness skin models consisting of multilayer skin equivalents were used for the investigation of MF effects on HSV-1 replication.
MF exhibited strong antiviral activity against HSV-1. The HSV-1-specific inhibitory concentration (IC50) was determined as 0.4μg/mL and the cytotoxic concentration (CC50) against Vero cells as 50μg/mL. A selectivity index (SI) (ratio of CC50 to IC50) of approximately 120 was calculated when MF was added to the virus inoculum for 1h at 37°C prior to infection. The replication of adenovirus 3 was not affected by MF.
MF abolished virus entry into the host cell by blocking viral attachment to the cell surface. When added after attachment at a concentration of >6μg/mL, the extract also inhibited penetration of HSV-1 into the host cell. Polyphenolic compounds from MF directly interacted with viral particles, leading to the oligomerisation of envelope proteins as demonstrated for the essential viral glycoprotein D (gD). Using organotypic full thickness tissue cultures, it was shown that treatment of HSV-1 infected cultures with the MF resulted in reduced viral spread.
A polyphenol-enriched extract from Myrothamnus flabellifolia strongly acts against HSV-1 by blocking viral entry into the cells.</abstract><cop>Shannon</cop><pub>Elsevier Ireland Ltd</pub><pmid>21211557</pmid><doi>10.1016/j.jep.2010.12.038</doi><tpages>7</tpages></addata></record> |
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| subjects | Adenoviridae - drug effects Adenoviridae Infections - microbiology Adenovirus Adsorption aerial parts Animals Antiviral Antiviral activity Antiviral Agents - pharmacology Antiviral Agents - therapeutic use antiviral properties Attachment Biological and medical sciences Cell culture Cell Line Cell surface Cercopithecus aethiops Cytotoxicity Envelope protein fluorescent antibody technique gD protein General pharmacology glycoprotein D glycoproteins Herpes Simplex - drug therapy Herpes Simplex - microbiology Herpes simplex virus 1 Herpes simplex virus type-1 Herpesvirus 1, Human - chemistry Herpesvirus 1, Human - drug effects Herpesvirus 1, Human - pathogenicity hosts Human herpesvirus 1 Humans Immunofluorescence Infection Inhibitory Concentration 50 Inoculum Keratinocytes - drug effects Keratinocytes - microbiology Magnoliopsida - chemistry mechanism of action Medical sciences Myrothamnaceae Myrothamnus Myrothamnus flabellifolia Welw Organotypic skin model Penetration Pharmacognosy. Homeopathy. Health food Pharmacology. Drug treatments Phytotherapy Plant Components, Aerial Plant diseases Plant Extracts - pharmacology Plant Extracts - therapeutic use Plaques Polyphenols Proanthocyanidins - pharmacology Proanthocyanidins - therapeutic use Replication Respiratory tract respiratory tract diseases Skin - drug effects Skin - microbiology Skin diseases Tissue culture Vero Cells Viral Envelope Proteins - chemistry virion Virus Integration - drug effects viruses Western blotting |
| title | Proanthocyanidin-enriched extract from Myrothamnus flabellifolia Welw. exerts antiviral activity against herpes simplex virus type 1 by inhibition of viral adsorption and penetration |
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