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http://www.elsevier.com/locate/toxicon/ An Interdisciplinary Journal on the Toxins Derived from Animals, Plants and Microorganisms. Official Journal of the International Society on Toxinology. Editorial Board Editor-in-Chief Alan L. Harvey, Strathclyde Institute for Drug Research, University of Strathclyde, 27 Taylor Street, Glasgow G4 0NR, UK Honorary Editors Philip Rosenberg, 37 Lincoln Avenue, West Orange, NJ 07052-2333, USA Review Editor Peter N. Strong, Sheffield Hallam University, Division of Biomedical Sciences, Sheffield, S1 1WB, UK Mini-Review Editors Ray S. Norton, Medicinal Chemistry and Drug Action, Monash Institute of Pharmaceutical Sciences, Faculty of Pharmacy and Pharmaceutical Sciences, Monash University (Parkville Campus), 381 Royal Parade, Parkville 3052, Australia Yara Cury, Laboratory of Physiopathology, Butantan Institute, Av. Vital Brazil, 1500, 05503-900 Sao Paulo, Brazil Classic toxins Review Editor Edward G. Rowan, Strathclyde Institute of Pharmacy and Biomedical Sciences, University of Strathclyde, Sir John Arbuthnott Building Room 301 F, 27 Taylor Street, Glasgow, G4 0NR, United Kingdom Editorial Council K. Aktories, Freiburg, Germany I. Asuzu, Nsukka, Nigeria G. L. Boyer, Syracuse, USA B. W. Brooks, Texas, USA J. J. Calvete, Valencia, Spain C. R. Carlini, Porto Alegre RS, Brazil E. E. Creppy, Bordeaux Cedex, France F. Ducancel, Gif-sur-Yvette, France M. Edery, Paris, France J. W. Fox, Charlottesville, VA, USA P. Gopalakrishnakone, Singapore J. M. Gutiérrez, San José, Costa Rica R. A. Harrison, Liverpool, UK W. C. Hodgson, Clayton, VIC, Australia R. J. Huxtable, Tucson, AZ, USA G. K. Isbister, Callaghan, NSW, Australia M. Ismail, Cairo, Egypt E. Kalapothakis, MG, Brazil W. R. Kem, Gainesville, FL, USA G. King, St. Lucia, QLD, Australia R. M. Kini, Singapore I. Krizaj, Ljubljana, Slovenia M. Lazdunski, Valbonne, France R. J. Lewis, Brisbane, QLD, Australia F. S. Markland Jr, Los Angeles, CA, USA M.-F. Martin-Eauclaire, Marseille, France D. Mebs, Frankfurt, Germany E. Moczydlowski, Albuquerque, NM, USA C. Montecucco, Padova, Italy B. A. Neilan, Sydney, NSW, Australia G. M. Nicholson, Sydney, NSW, Australia B. M. Olivera, Salt Lake City, UT, USA S. Pflugmacher, Berlin, Germany M. A. Poli, Maryland, USA L. D. Possani, Cuernavaca, Mexico S. M. T. Serrano, Sao Paulo, Brazil W. T. Shier, Minneapolis, MN, USA K. Sivonen, Helsinki, Finland T. Tamiya, Tokyo, Japan A. Tubaro, Trieste, Italy J. Tytgat, Leuven, Belgium D. A. Warrell, Oxford, UK J. White, North Adelaide, Australia Y. Zhang, Yunnan, China R. B. Zingali, Rio de Janeiro, Brazil Printed by Henry Ling Ltd, The Dorset Press, Dorchester, UK International Society on Toxinology President: Prof. Alan L. Harvey, Strathclyde Institute for Drug Research, University of Strathclyde, 27 Taylor Street, Glasgow G4 0NR, UK Secretary-Treasurer: Prof. Julian White, c/o Toxinology Dept., Women’s & Children’s Hospital, North Adelaide SA 5006, AUSTRALIA The purpose of this society is to advance knowledge on the properties of toxins and antitoxins and to bring together scholars interested in these substances through a common society. Membership consists of those who have conducted and published meritorious original investigations in toxinology, while persons who do not qualify for membership but are interested in the field of toxinology are eligible for associate membership. Further information concerning the Society and applications for membership may be obtained from the Secretary-Treasurer. Publication information: Toxicon (ISSN 0041-0101) For 2014, volumes 77–92 (16 issues) are scheduled for publication. Subscription prices are available upon request from the Publisher or from the Elsevier Customer Service Department nearest you or from this journal’s website (http://www.elsevier.com/locate/toxicon). Further information is available on this journal and other Elsevier products through Elsevier’s website (http://www.elsevier.com). Subscriptions are accepted on a prepaid basis only and are entered on a calendar year basis. Issues are sent by standard mail (surface within Europe, air delivery outside Europe). Priority rates are available upon request. Claims for missing issues should be made within six months of the date of dispatch. Orders, claims, and journal inquiries: please contact the Elsevier Customer Service Department nearest you: St. Louis: Elsevier Customer Service Department, 3251 Riverport Lane, Maryland Heights, MO 63043, USA; phone: (877) 8397126 [toll free within the USA]; (+1) (314) 4478878 [outside the USA]; fax: (+1) (314) 4478077; e-mail: [email protected] Oxford: Elsevier Customer Service Department, The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, UK; phone: (+44) (1865) 843434; fax: (+44) (1865) 843970; e-mail: [email protected] Tokyo: Elsevier Customer Service Department, 4F Higashi-Azabu, 1-Chome Bldg, 1-9-15 Higashi-Azabu, Minato-ku, Tokyo 106-0044, Japan; phone: (+81) (3) 5561 5037; fax: (+81) (3) 5561 5047; e-mail: [email protected] Singapore: Elsevier Customer Service Department, 3 Killiney Road, #08-01 Winsland House I, Singapore 239519; phone: (+65) 6349 0222; fax: (+65) 6733 1510; e-mail: [email protected] Author inquiries: For inquiries relating to the submission of articles (including electronic submission) please visit this journal’s homepage at http://www.elsevier.com/locate/toxicon. For detailed instructions on the preparation of electronic artwork, please visit http://www.elsevier.com/artworkinstructions. Contact details for questions arising after acceptance of an article, especially those relating to proofs, will be provided by the publisher. You can track accepted articles at http://www.elsevier.com/trackarticle. You can also check our Author FAQs at http://www.elsevier.com/ authorFAQ and/or contact Customer Support via http://support.elsevier.com. Sponsored Supplements and/or Commercial Reprints: For more information please contact Elsevier Life Sciences Commercial Sales, Radarweg 29, 1043 NX Amsterdam, The Netherlands; Phone: (+31) (20) 485 2939 / 2059; e-mail: [email protected]. Advertising information. If you are interested in advertising or other commercial opportunities please e-mail: [email protected] and your inquiry will be passed to the correct person who will respond to you within 48 hours. USA mailing notice: Toxicon (ISSN 0041-0101) is published (sixteen issues per year, one per month with extra issues in March, June, September and December) by Elsevier Ltd (The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, UK). Periodicals postage paid at Jamaica, NY 11431 and additional mailing offices. USA POSTMASTER: Send change of address to Toxicon, Elsevier Customer Service Department, 3251 Riverport Lane, Maryland Heights, MO 63043, USA. AIRFREIGHT AND MAILING in USA by Air Business Ltd., c/o Worldnet Shipping Inc., 156-15, 146th Avenue, 2nd Floor, Jamaica, NY 11434, USA. Cover design by Xavier Sprungli from the Toxinomics Foundation, Geneva, Switzerland (www.toxinomics.org). 3D molecule is the mu-conopeptide CnIIIC (XEP-018) from the cone snail Conus consors (Favreau et al., Brit. J. Pharmacol., 2012) realised by Henry Hocking, Ludovic Carlier and Rolf Boelens, University of Utrecht, The Netherlands (www. uu.nl). The Conus consors harpoon (radula) picture is from Jean-Jacques Soin, France. The Dendroaspis viridis (Western green mamba) snake picture is by Reto Stöcklin from the Atheris Laboratories, Geneva, Switzerland (www.atheris.com). We are most grateful to the Governement of New Caledonia, to Reptiles du Monde (Servion, Switzerland), to the French Institut de Recherche pour le Développement (IRD) and, last but not least, to the European Commission for its support and funding of “CONCO, the cone snail genome project for health” (LSHB-CT-2007-037592; www.conco.eu) with Dr. Torbjörn Ingemansson as principal scientific officer. © 2014 The Toxinomics Foundation. The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper) For a full and complete Guide for Authors, please go to: http://www.elsevier.com/locate/toxicon/

Editorial Board/Publication Information

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http://www.elsevier.com/locate/toxicon/An Interdisciplinary Journal on the Toxins Derived from Animals, Plants and Microorganisms.

Official Journal of the International Society on Toxinology.

Editorial BoardEditor-in-Chief

Alan L. Harvey, Strathclyde Institute for Drug Research, University of Strathclyde, 27 Taylor Street, Glasgow G4 0NR, UK

Honorary EditorsPhilip Rosenberg, 37 Lincoln Avenue, West Orange, NJ 07052-2333, USA

Review Editor

Peter N. Strong, Sheffield Hallam University, Division of Biomedical Sciences, Sheffield, S1 1WB, UK

Mini-Review EditorsRay S. Norton, Medicinal Chemistry and Drug Action, Monash Institute of Pharmaceutical Sciences, Faculty of Pharmacy

and Pharmaceutical Sciences, Monash University (Parkville Campus), 381 Royal Parade, Parkville 3052, Australia Yara Cury, Laboratory of Physiopathology, Butantan Institute, Av. Vital Brazil, 1500, 05503-900 Sao Paulo, Brazil

Classic toxins Review Editor Edward G. Rowan, Strathclyde Institute of Pharmacy and Biomedical Sciences, University of Strathclyde,

Sir John Arbuthnott Building Room 301 F, 27 Taylor Street, Glasgow, G4 0NR, United Kingdom

Editorial Council

K. Aktories, Freiburg, GermanyI. Asuzu, Nsukka, NigeriaG. L. Boyer, Syracuse, USAB. W. Brooks, Texas, USAJ. J. Calvete, Valencia, SpainC. R. Carlini, Porto Alegre RS, BrazilE. E. Creppy, Bordeaux Cedex, FranceF. Ducancel, Gif-sur-Yvette, FranceM. Edery, Paris, FranceJ. W. Fox, Charlottesville, VA, USAP. Gopalakrishnakone, SingaporeJ. M. Gutiérrez, San José, Costa RicaR. A. Harrison, Liverpool, UKW. C. Hodgson, Clayton, VIC, AustraliaR. J. Huxtable, Tucson, AZ, USA

G. K. Isbister, Callaghan, NSW, AustraliaM. Ismail, Cairo, EgyptE. Kalapothakis, MG, BrazilW. R. Kem, Gainesville, FL, USAG. King, St. Lucia, QLD, AustraliaR. M. Kini, SingaporeI. Krizaj, Ljubljana, SloveniaM. Lazdunski, Valbonne, FranceR. J. Lewis, Brisbane, QLD, AustraliaF. S. Markland Jr, Los Angeles, CA, USAM.-F. Martin-Eauclaire, Marseille, FranceD. Mebs, Frankfurt, GermanyE. Moczydlowski, Albuquerque, NM, USAC. Montecucco, Padova, ItalyB. A. Neilan, Sydney, NSW, Australia

G. M. Nicholson, Sydney, NSW, AustraliaB. M. Olivera, Salt Lake City, UT, USAS. Pflugmacher, Berlin, GermanyM. A. Poli, Maryland, USAL. D. Possani, Cuernavaca, MexicoS. M. T. Serrano, Sao Paulo, BrazilW. T. Shier, Minneapolis, MN, USAK. Sivonen, Helsinki, FinlandT. Tamiya, Tokyo, JapanA. Tubaro, Trieste, ItalyJ. Tytgat, Leuven, BelgiumD. A. Warrell, Oxford, UKJ. White, North Adelaide, AustraliaY. Zhang, Yunnan, ChinaR. B. Zingali, Rio de Janeiro, Brazil

Printed by Henry Ling Ltd, The Dorset Press, Dorchester, UK

International Society on ToxinologyPresident: Prof. Alan L. Harvey, Strathclyde Institute for Drug Research, University of Strathclyde, 27 Taylor Street, Glasgow G4 0NR, UKSecretary-Treasurer: Prof. Julian White, c/o Toxinology Dept., Women’s & Children’s Hospital, North Adelaide SA 5006, AUSTRALIA The purpose of this society is to advance knowledge on the properties of toxins and antitoxins and to bring together scholars interested in these substances through a common society. Membership consists of those who have conducted and published meritorious original investigations in toxinology, while persons who do not qualify for membership but are interested in the field of toxinology are eligible for associate membership. Further information concerning the Society and applications for membership may be obtained from the Secretary-Treasurer.

Publication information: Toxicon (ISSN 0041-0101) For 2014, volumes 77–92 (16 issues) are scheduled for publication. Subscription prices are available upon request from the Publisher or from the Elsevier Customer Service Department nearest you or from this journal’s website (http://www.elsevier.com/locate/toxicon). Further information is available on this journal and other Elsevier products through Elsevier’s website (http://www.elsevier.com). Subscriptions are accepted on a prepaid basis only and are entered on a calendar year basis. Issues are sent by standard mail (surface within Europe, air delivery outside Europe). Priority rates are available upon request. Claims for missing issues should be made within six months of the date of dispatch. Orders, claims, and journal inquiries: please contact the Elsevier Customer Service Department nearest you:St. Louis: Elsevier Customer Service Department, 3251 Riverport Lane, Maryland Heights, MO 63043, USA; phone: (877) 8397126 [toll free within the USA]; (+1) (314) 4478878 [outside the USA]; fax: (+1) (314) 4478077; e-mail: [email protected]: Elsevier Customer Service Department, The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, UK; phone: (+44) (1865) 843434; fax: (+44) (1865) 843970; e-mail: [email protected] Tokyo: Elsevier Customer Service Department, 4F Higashi-Azabu, 1-Chome Bldg, 1-9-15 Higashi-Azabu, Minato-ku, Tokyo 106-0044, Japan; phone: (+81) (3) 5561 5037; fax: (+81) (3) 5561 5047; e-mail: [email protected] Singapore: Elsevier Customer Service Department, 3 Killiney Road, #08-01 Winsland House I, Singapore 239519; phone: (+65) 6349 0222; fax: (+65) 6733 1510; e-mail: [email protected] Author inquiries: For inquiries relating to the submission of articles (including electronic submission) please visit this journal’s homepage at http://www.elsevier.com/locate/toxicon. For detailed instructions on the preparation of electronic artwork, please visit http://www.elsevier.com/artworkinstructions. Contact details for questions arising after acceptance of an article, especially those relating to proofs, will be provided by the publisher. You can track accepted articles at http://www.elsevier.com/trackarticle. You can also check our Author FAQs at http://www.elsevier.com/authorFAQ and/or contact Customer Support via http://support.elsevier.com.Sponsored Supplements and/or Commercial Reprints: For more information please contact Elsevier Life Sciences Commercial Sales, Radarweg 29, 1043 NX Amsterdam, The Netherlands; Phone: (+31) (20) 485 2939 / 2059; e-mail: [email protected].

Advertising information. If you are interested in advertising or other commercial opportunities please e-mail: [email protected] and your inquiry will be passed to the correct person who will respond to you within 48 hours. USA mailing notice: Toxicon (ISSN 0041-0101) is published (sixteen issues per year, one per month with extra issues in March, June, September and December) by Elsevier Ltd (The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, UK). Periodicals postage paid at Jamaica, NY 11431 and additional mailing offices.USA POSTMASTER: Send change of address to Toxicon, Elsevier Customer Service Department, 3251 Riverport Lane, Maryland Heights, MO 63043, USA.AIRFREIGHT AND MAILING in USA by Air Business Ltd., c/o Worldnet Shipping Inc., 156-15, 146th Avenue, 2nd Floor, Jamaica, NY 11434, USA.

Cover design by Xavier Sprungli from the Toxinomics Foundation, Geneva, Switzerland (www.toxinomics.org). 3D molecule is the mu-conopeptide CnIIIC (XEP-018) from the cone snail Conus consors (Favreau et al., Brit. J. Pharmacol., 2012) realised by Henry Hocking, Ludovic Carlier and Rolf Boelens, University of Utrecht, The Netherlands (www.uu.nl). The Conus consors harpoon (radula) picture is from Jean-Jacques Soin, France. The Dendroaspis viridis (Western green mamba) snake picture is by Reto Stöcklin from the Atheris Laboratories, Geneva, Switzerland (www.atheris.com). We are most grateful to the Governement of New Caledonia, to Reptiles du Monde (Servion, Switzerland), to the French Institut de Recherche pour le Développement (IRD) and, last but not least, to the European Commission for its support and funding of “CONCO, the cone snail genome project for health” (LSHB-CT-2007-037592; www.conco.eu) with Dr. Torbjörn Ingemansson as principal scientific officer. © 2014 The Toxinomics Foundation.

∞ The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper)

For a full and complete Guide for Authors, please go to: http://www.elsevier.com/locate/toxicon/