Handbook of Venoms and Toxins of Reptiles, 2nd Edition

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Contents
Preface.......................................................................................................................................................................................... ix
About the Editor............................................................................................................................................................................ xi
Contributors................................................................................................................................................................................xiii

SECTION I Introduction and Technologies Used in Toxinology

Chapter 1 Reptile Venoms and Toxins: Unlimited Opportunities for Basic and Applied Research........................................ 3
Stephen P. Mackessy

Chapter 2 Present and Future of Snake Venom Proteomics Profiling.................................................................................... 19

Juan J. Calvete and Bruno Lomonte

Chapter 3 Applications of Genomics and Related Technologies for Studying Reptile Venoms............................................ 29
Drew R. Schield, Blair W. Perry, Giulia I.M. Pasquesi, Richard W. Orton, Zachary L. Nikolakis,
Aundrea K. Westfall and Todd A. Castoe

Chapter 4 Snake Venom Gland Transcriptomics.................................................................................................................... 43

Cassandra M. Modahl and Rajeev Kungur Brahma

Chapter 5 X-ray Crystallography and Structural Studies of Toxins....................................................................................... 59

Vinícius Lucatelle da Silva, Ricardo Barros Mariutti, Mônika Aparecida Coronado, Raphael Josef
Eberle, Fábio Rogério de Moraes and Raghuvir Krishnaswamy Arni

Chapter 6 Envenomations and Treatment: Translating between the Bench and the Bedside................................................. 73
Nicklaus Brandehoff and Jordan Benjamin

Chapter 7 Current Assessment of the State of Snake Venom Toxinological Research with a View to the Future................ 79
Sarah Natalie Cirilo Gimenes and Jay W. Fox

SECTION II Venom Gland Structure, Systematics and Ecology

Chapter 8 Reptile Venom Glands: Form, Function, Future, Concepts and Controversies..................................................... 99
Scott A. Weinstein

Chapter 9 Advances in Venomous Snake Systematics, 2009–2019..................................................................................... 123

Wolfgang Wüster

Chapter 10 Biochemical Ecology of Venomous Snakes..........................................................................................................147

Cara F. Smith and Stephen P. Mackessy

v
vi Contents

Chapter 11 Resistance of Native Species to Reptile Venoms..................................................................................................161

Danielle H. Drabeck

SECTION III Reptile Venom Non-Enzymatic Toxins

Chapter 12 Three-Finger Toxins............................................................................................................................................. 177

Rajeev Kungur Brahma, Cassandra M. Modahl and R. Manjunatha Kini

Chapter 13 Myotoxin a, Crotamine and Defensin Homologs in Reptile Venoms.................................................................. 195

Lucas C. Porta, Pedro Z. Amaral, Paulo Z. Amaral and Mirian A. F. Hayashi

Chapter 14 Reptile Venom Disintegrins..................................................................................................................................211

Anthony J. Saviola and Juan J. Calvete

Chapter 15 Reptile Venom Cysteine-Rich Secretory Proteins............................................................................................... 225

María Elisa Peichoto and Marcelo Larami Santoro

Chapter 16 Bradykinin-Potentiating and Related Peptides from Reptile Venoms................................................................. 241

Daniel Carvalho Pimenta and Patrick Jack Spencer

Chapter 17 Exendin-4 and Its Related Peptides...................................................................................................................... 251

Michelle Khai Khun Yap and Nurhamimah Misuan

Chapter 18 Reptile Venom C-Type Lectins............................................................................................................................ 271

Kenneth J. Clemetson

Chapter 19 Snake Venom Kunitz-type Inhibitors and Cystatins – Structure and Function................................................... 285
Elda E. Sánchez, Emelyn Salazar, Montamas Suntravat and Francisco Torres

Chapter 20 Small Molecular Constituents of Snake Venoms................................................................................................. 305

Alejandro Villar-Briones and Steven D. Aird

Chapter 21 Cobra Venom Factor: Structure, Function, Biology, Research Tool, and Drug Lead.......................................... 323
Carl-Wilhelm Vogel, Brian E. Hew and David C. Fritzinger

Chapter 22 Snake Toxins Targeting Diverse Ion Channels.................................................................................................... 339

Matan Geron and Avi Priel

SECTION IV Reptile Venom Enzyme Toxins

Chapter 23 Thrombin-Like Serine Proteinases in Reptile Venoms .......................................................................................351

Stephen D. Swenson, Samantha Stack and Francis S. Markland Jr.
Contents vii

Chapter 24 Snake Venom Metalloproteinases........................................................................................................................ 363

Charlotte A. Dawson, Stuart Ainsworth, Laura-Oana Albulescu and Nicholas R. Casewell

Chapter 25 Snake Venom Matrix Metalloproteinases (svMMPs): Alternative Proteolytic Enzymes in Rear-Fanged
Snake Venoms...................................................................................................................................................... 381
Inácio L. M. Junqueira-de-Azevedo and Juan David Bayona-Serrano

Chapter 26 Snake Venom Phospholipase A2 Toxins............................................................................................................... 389

Bruno Lomonte and Igor Križaj

Chapter 27 Reptile Venom L-Amino Acid Oxidases – Structure and Function.....................................................................413

Juliana P. Zuliani, Mauro V. Paloschi, Adriana S. Pontes, Charles N. Boeno, Jéssica A. Lopes,
Sulamita S. Setubal, Fernando B. Zanchi and Andreimar M. Soares

Chapter 28 Snake Venom Nucleases, Nucleotidases and Phosphomonoesterases..................................................................431

Jüri Siigur and Ene Siigur

Chapter 29 Reptile Venom Acetylcholinesterases.................................................................................................................. 445

Mushtaq Ahmed, Wasim Ahmad, Nadia Mushtaq, Rehmat Ali Khan and Maria Rosa Chitolina Schetinger

Chapter 30 Inhibitors of Reptile Venom Toxins..................................................................................................................... 453

Ana F. Gómez Garay, Jorge J. Alfonso, Anderson M. Kayano, Juliana C. Sobrinho,
Cleopatra A. S. Caldeira, Rafaela Diniz-Sousa, Fernando B. Zanchi, Andreimar M. Soares and
Juliana P. Zuliani

SECTION V Global Approaches to Envenomations and Treatments

Chapter 31 Snakebite Envenomation as a Neglected Tropical Disease: New Impetus for Confronting an Old Scourge...... 471
José María Gutiérrez

Chapter 32 Current Industrial Production of Snake Antivenoms........................................................................................... 485

Mariángela Vargas, Melvin Sánchez, Andrés Hernández, Aarón Gómez, Mauricio Arguedas,
Andrés Sánchez, Laura Sánchez, Mauren Villalta, María Herrera and Álvaro Segura

Chapter 33 Antivenom in the Age of Recombinant DNA Technology.................................................................................. 499

Andreas H. Laustsen

Chapter 34 Epidemiology and Treatment of Reptile Envenomations in the United States.....................................................511

Daniel E. Keyler and Nicklaus Brandehoff

Chapter 35 Envenomations by Reptiles in Mexico................................................................................................................. 529

Edgar Neri-Castro, Melisa Bénard-Valle, Jorge López de León,
Leslie Boyer and Alejandro Alagón
viii Contents

Chapter 36 Snakebite Envenomation in Central America: Epidemiology, Pathophysiology and Treatment......................... 543
José María Gutiérrez

Chapter 37 Snakebite in Southeast Asia: Envenomation and Clinical Management............................................................. 559

Nget Hong Tan, Kae Yi Tan and Choo Hock Tan

Chapter 38 Snake Envenomation: Therapy and Challenges in India..................................................................................... 581

Ashis K. Mukherjee, Bhargab Kalita, Sumita Dutta, Aparup Patra, Chitta R. Maiti and Dileep Punde

Chapter 39 Snakebite in Africa: Current Situation and Urgent Needs................................................................................... 593

Jean-Philippe Chippaux

Chapter 40 Approaches to Snake Envenomation in Southern Africa......................................................................................613

James Pattinson, George Oosthuizen, Colin R. Tilbury and Darryl Wood

SECTION VI Reptile Venoms – Production and as a Source of Therapeutics

Chapter 41 Large-Scale Snake Colonies for Venom Production: Considerations and Challenges........................................ 623
Kristen L. Wiley and James R. Harrison

Chapter 42 Toxins to Drugs – Biochemistry and Pharmacology........................................................................................... 635

Zoltan Takacs
Index.......................................................................................................................................................................................... 653
Preface
It has been 10 years since the first edition of the Handbook This second edition is being assembled under one of the
was published, and the field of toxinology has changed exten- most unusual global health crises to arise in modern times,
sively. This second edition represents a greatly expanded ver- certainly in my lifetime – the SARS-CoV-2 virus and the
sion of the first edition, with an attempt to capture much of the Covid-19 pandemic, currently (January 17, 2021) afflicting
recent developments in the field, and most of it is new mate- nearly 95 million people globally. In spite of this, and the
rial, with many new authors. The format is similar to the first many challenges that this pandemic has caused for everyone,
edition, but the information contained in this second edition the authors of the chapters contained herein have been respon-
is a complement to the first rather than a replacement. The sive and timely in their submissions, and I greatly appreciate
six sections of this edition present current information from and thank them for their efforts.
many of the leading researchers and physicians in toxinology In the 10 years since the last volume, many of the most
(defined broadly), and topics range from functional morphol- prominent names in toxinology and herpetology have passed,
ogy, evolution and ecology to biochemistry, crystallography, including Findlay E. Russell, David Chiszar, Hobart M. Smith,
-omics technologies and more. With the recent recognition Kenneth V. Kardong and most recently, Alan L. Harvey. I
(again) by the World Health Organization of snakebite as a had the great fortune to interact with all these fine scientists,
neglected tropical disease, there has been renewed effort by have benefited tremendously from these experiences and am
many agencies, including non-governmental organizations grateful to have had the opportunity to know them. Their con-
and governmental groups, to address this scourge, which tributions to the field are extensive, the impact of their work
differentially afflicts those least able to afford the necessary remains significant and timely, and they are greatly missed.
treatment. To this end, the section on snakebite has been I would like to express particular gratitude to Ken Kardong,
expanded and includes several chapters dealing with the my PhD advisor and mentor, for his patience and guidance
problem broadly and with new technologies and the prom- during my years at Washington State University, and to David
ises these new approaches may hold to counter the deleterious Chiszar for his advice and support at an early point in my
effects of envenomation. career and for continuing collaborations.
Shortly after the first edition was published, I attended No projects exist in a vacuum, and this book is no excep-
the European section meeting of the International Society on tion. I thank my many students, graduate and undergraduate,
Toxinology, held in Valencia, Spain and hosted by Dr. Juan for their excellent efforts in research, which have produced
Calvete and colleagues. As a regular attendee of International many fine papers, and I congratulate them on their many
Society on Toxinology conferences, I was aware of the general successes. In particular, I would like to acknowledge the
rate of progress in toxinology, which had increased in recent outstanding work of Dr. Anthony Saviola, Dr. Cassandra
years, but I was unprepared for the exponential increase in Modahl and PhD candidate Cara Smith – your work is inspi-
level of technical complexity, detail and sophistication seen rational and has contributed greatly to the success of our lab.
at the presentations given in Valencia. I remember think- Collaborations with visiting and other scientists, including Dr.
ing “Well, maybe it’s time to hang it up, move to a different Ashis Mukherjee and Dr. María Elisa Peichoto, have greatly
field, before I’m swept away by this tidal wave of progress”. enhanced my lab and the experiences for my students, and
However, I did not leave the field, and it has been very satisfy- ongoing collaborative work with Dr. Todd A. Castoe and his
ing to see how toxinology, particularly those areas concerned group has opened up many new research directions involving
with analyses of reptile venoms, has developed and matured venomous snake genomics. Finally, I would like to recognize
since that time. I have also had the good fortune to collaborate the love, help, support and encouragement of my life partner,
with many excellent scientists on a variety of topics dealing Dr. Debra Kaye Holman, and the love and encouragement of
with venomous snakes and their venoms, and it is through my daughter, Elizabeth K. Mackessy; I dedicate this book to
collaborations like these, with individuals having expertise in Kaye and Elizabeth, and to my brothers and sisters, Denise,
many different areas, that even more rapid progress can occur. Kristine, Tom, John and Eileen.

ix
About the Editor
Stephen P. Mackessy is Professor of Biology in the School of Dr. Mackessy also teaches graduate and undergradu-
Biological Sciences at the University of Northern Colorado. ate courses in biomedicine (Toxinology of Snake Venoms,
His research broadly encompasses the biology of venom- Current Topics in Biomedical Research, Parasitology,
ous snakes and the biochemistry of snake venoms, and he Human Anatomy) and vertebrate biology (Herpetology,
has published over 180 scientific papers, book chapters and Comparative Anatomy) at the University of Northern
natural history notes, several books including Handbook of Colorado, where he has received awards in recognition of
Venoms and Toxins of Reptiles – CRC Press and The Biology outstanding research and teaching (1999 – Distinguished
of Rattlesnakes II – ECO Herpetological Publishing, and Scholar Award; 2004 – The Joseph Lazlo Memorial Award
special editions of the Journal of Toxicology-Toxin Reviews, for research – International Herpetological Society; 2006–7
Toxicon and Toxins. His research has included many gradu- – NHS Excellence in Scholarship; 2011–12 – NHS Faculty
ate and undergraduate students as well as collaborations Mentor of the Year – Graduate; 2012 – M. Lucille Harrison
with colleagues from Singapore, Spain, México, Costa Rica, Award (the University of Northern Colorado’s top fac-
Argentina, Brazil, France, India, University of Texas and vari- ulty honor); 2020 – Meritorious Teaching and Mentoring
ous other universities in the United States. Several ongoing Award – Herpetology – Joint Meeting of Ichthyologists and
projects are centered on understanding the evolution and reg- Herpetologists). He earned a BA and an MA in Biology
ulation of venom systems in snakes and the biological signifi- (Ecology and Evolution section) at the University of
cance of venom compositional variation, particularly in areas California at Santa Barbara, Department of Biology (with
of introgression, with an overarching interest in the interface Dr. S.S. Sweet), and his PhD (with a minor in Biochemistry)
of snake ecology/evolution and venom biochemistry/phar- was received from Washington State University, Department
macology. Broad sampling of venoms from many species of of Zoology (with Dr. K.V. Kardong). He was a postdoctoral
rattlesnakes (Crotalus, Sistrurus), other vipers, seasnakes and research associate at Colorado State University, Department
numerous species of rear-fanged snakes has resulted in exten- of Biochemistry and Molecular Biology (with Dr. A.T. Tu)
sive fieldwork in the western United States, Mexico, Costa before joining the School of Biological Sciences at the
Rica, Guam, Taiwan and Southeast Asia. Other projects have University of Northern Colorado. He was the managing edi-
focused on the effects of venoms and toxins on metastatic tor of the Journal of Natural Toxins for 7 years and he has
cell proliferation and the investigation of novel toxins for new served as a scientific peer reviewer for over 70 different jour-
drug leads, structure/function relationships among venom nals. His research has been featured in films by the BBC and
toxin families and, more recently, the application of genomic Discovery Channel and has been the subject of many media
and proteomic approaches toward understanding venom bio- publications. Personal interests include fieldwork with ven-
chemistry, pharmacology and evolution. His research pro- omous snakes, music and motorcycles, as well as traveling
gram has been supported by many local, state and national and camping.
funding agencies.

xi
Contributors
Wasim Ahmad Mauricio Arguedas
Department of Biotechnology Instituto Clodomiro Picado
University of Science and Technology Universidad de Costa Rica
Bannu, Pakistan San José, Costa Rica

Mushtaq Ahmed Raghuvir Krishnaswamy Arni

Department of Biotechnology Structural Biology Group
University of Science and Technology Department of Physics, IBILCE/UNESP
Bannu, Pakistan Multiuser Center for Biomolecular
Innovation
Stuart Ainsworth São José do Rio Preto, Brazil
Centre for Snakebite Research & Interventions
Liverpool School of Tropical Medicine
Juan David Bayona-Serrano
Liverpool, United Kingdom
Laboratório de Toxinologia Aplicada Instituto
Butantan
Steven D. Aird
São Paulo, Brazil
Ecology and Evolution Unit
Okinawa Institute of Science and Technology
Okinawa-ken, Japan Melisa Bénard-Valle
Technical Editor, Japan Instituto de Biotecnología
Universidad Nacional Autónoma de México
Alejandro Alagón Cuernavaca, México
Instituto de Biotecnología
Universidad Nacional Autónoma de México Jordan Benjamin
Cuernavaca, México Asclepius Snakebite Foundation
Seattle, Washington
Laura-Oana Albulescu and
Centre for Snakebite Research & Interventions Whitman College
Liverpool School of Tropical Medicine Walla Walla, Washington
Liverpool, United Kingdom
Charles N. Boeno
Jorge J. Alfonso Laboratório de Imunologia Aplicada à Saúde Fundação
LaBioProt, CEBio, Fundação Oswaldo Cruz Oswaldo Cruz
Unidade Rondônia and Departamento de Medicina Unidade Rondônia
Universidade Federal de Rondônia Porto Velho, Brazil
Porto Velho, Brazil
and
Leslie Boyer
Centro para el Desarrollo de la Investigación Científica
Venom Immunochemistry, Pharmacology and Emergency
Asunción, Paraguay
Response Institute
University of Arizona
Paulo Z. Amaral
Tucson, Arizona
Department of Pharmacology
Escola Paulista de Medicina
Universidade Federal de São Paulo Rajeev Kungur Brahma
São Paulo, Brazil Protein Science Lab
Department of Biological Sciences
Pedro Z. Amaral University of Singapore, Singapore
Department of Pharmacology
Escola Paulista de Medicina Nicklaus Brandehoff
Universidade Federal de São Paulo Rocky Mountain Poison and Drug Safety
São Paulo, Brazil Denver, Colorado

xiii
xiv Contributors

Cleopatra A. S. Caldeira Sumita Dutta

BioProt, CEBio, Fundação Oswaldo Cruz Microbial Biotechnology and Protein Research Laboratory
Unidade Rondônia and Departamento de Medicina Department of Molecular Biology and Biotechnology
Universidade Federal de Rondônia School of Sciences
Porto Velho, Brazil Tezpur University
Tezpur, India
Juan J. Calvete
Laboratorio de Venómica Evolutiva y Traslacional Raphael Josef Eberle
Instituto de Biomedicina de Valencia C.S.I.C. Structural Biology Group
Valencia, Spain Multiuser Center for Biomolecular Innovation
Department of Physics, IBILCE/UNESP
Nicholas R. Casewell São José do Rio Preto, Brazil
Centre for Snakebite Research & Interventions
Liverpool School of Tropical Medicine
Jay W. Fox
Liverpool, United Kingdom
University of Virginia School of Medicine
Charlottesville, Virginia
Todd A. Castoe
Department of Biology
University of Texas at Arlington David C. Fritzinger
Arlington, Texas University of Hawaii Cancer Center
Honolulu, Hawaii
Jean-Philippe Chippaux
Université de Paris, MERIT, IRD, France Matan Geron
Institut Pasteur, CRT The Institute for Drug Research
Paris, France School of Pharmacy
The Hebrew University of Jerusalem
Kenneth J. Clemetson Jerusalem, Israel
Theodor Kocher Institute
University of Bern Sarah Natalie Cirilo Gimenes
Bern, Switzerland Butantan Institute
São Paulo, Brazil
Mônika Aparecida Coronado
Structural Biology Group
Aarón Gómez
Multiuser Center for Biomolecular Innovation
Instituto Clodomiro Picado
Department of Physics, IBILCE/UNESP
Universidad de Costa Rica
São José do Rio Preto, Brazil
San José, Costa Rica
Charlotte A. Dawson
Centre for Snakebite Research & Interventions Ana F. Gómez Garay
Liverpool School of Tropical Medicine LaBioProt, CEBio, Fundação Oswaldo Cruz
Liverpool, United Kingdom Unidade Rondônia and Departamento de Medicina
Universidade Federal de Rondônia
Rafaela Diniz-Sousa Porto Velho, Brazil
LaBioProt, CEBio, Fundação Oswaldo Cruz and
Unidade Rondônia and Departamento de Centro para el Desarrollo de la Investigación Científica
Medicina Asunción, Paraguay
Universidade Federal de Rondônia
and José María Gutiérrez
Centro Universitário São Lucas Instituto Clodomiro Picado
Porto Velho, Brazil Universidad de Costa Rica
San José, Costa Rica
Danielle H. Drabeck
Department of Ecology, Evolution and Behavior James R. Harrison
University of Minnesota Kentucky Reptile Zoo
St. Paul, Minnesota Slade, Kentucky
Contributors xv

Mirian A. F. Hayashi R. Manjunatha Kini

Department of Pharmacology Protein Science Lab
Escola Paulista de Medicina Department of Biological Sciences
Universidade Federal de São Paulo University of Singapore, Singapore
São Paulo, Brazil
and Igor Križaj
National Institute for Translational Medicine (INCT-TM, Jožef Stefan Institute
CNPq/FAPESP/CAPES) Department of Molecular and Biomedical
Brazil Sciences
Ljubljana, Slovenia
Andrés Hernández
Instituto Clodomiro Picado Andreas H. Laustsen
Universidad de Costa Rica Department of Biotechnology and Biomedicine
San José, Costa Rica Technical University of Denmark
Kongens Lyngby, Denmark
María Herrera
Instituto Clodomiro Picado Bruno Lomonte
Universidad de Costa Rica Instituto Clodomiro Picado
San José, Costa Rica Universidad de Costa Rica
San José, Costa Rica
Brian E. Hew
University of Hawaii Cancer Center Jéssica A. Lopes
Honolulu, Hawaii Laboratório de Imunologia Aplicada à Saúde Fundação
Oswaldo Cruz
Inácio L. M. Junqueira-de-Azevedo Unidade Rondônia
Laboratório de Toxinologia Aplicada Instituto Butantan Porto Velho, Brazil
São Paulo, Brazil
Jorge López de León
Bhargab Kalita Hospital General Norberto Treviño Zapata Ciudad
Microbial Biotechnology and Protein Research Victoria
Laboratory Tamaulipas, México
Department of Molecular Biology
and Stephen P. Mackessy
Biotechnology, School of Sciences School of Biological Sciences
Tezpur University University of Northern Colorado
Tezpur, India Greeley, Colorado

Anderson M. Kayano Chitta R. Maiti

BioProt, CEBio, Fundação Oswaldo Cruz Department of Biochemistry
Unidade Rondônia and Departamento de Medicina Burdwan Medical College
Universidade Federal de Rondônia Burdwan, India
and
Centro de Pesquisa em Medicina Tropical Ricardo Barros Mariutti
Porto Velho, Brazil Structural Biology Group
Multiuser Center for Biomolecular Innovation
Daniel E. Keyler Department of Physics, IBILCE/UNESP
Department of Experimental and Clinical Pharmacology São José do Rio Preto, Brazil
University of Minnesota
Minneapolis, Minnesota Francis S. Markland Jr.
Department of Biochemistry and Molecular
Rehmat Ali Khan Medicine
Department of Biotechnology Keck School of Medicine
University of Science and Technology University of Southern California
Bannu, Pakistan Los Angeles, California
xvi Contributors

Nurhamimah Misuan Giulia I. M. Pasquesi

School of Science Department of Biology
Monash University Malaysia University of Texas at Arlington
Selangor, Malaysia Arlington, Texas

Cassandra M. Modahl Aparup Patra

Protein Science Lab Microbial Biotechnology and Protein Research Laboratory
Department of Biological Sciences Department of Molecular Biology and Biotechnology
University of Singapore, Singapore School of Sciences
Tezpur University
Fábio Rogério de Moraes Tezpur, India
Structural Biology Group
Multiuser Center for Biomolecular Innovation James Pattinson
Department of Physics, IBILCE/UNESP Department of Surgery
São José do Rio Preto, Brazil Chris Hani Baragwanath Academic Hospital
Gauteng, South Africa
Ashis K. Mukherjee
Microbial Biotechnology and Protein Research Laboratory María Elisa Peichoto
Department of Molecular Biology and Biotechnology Consejo Nacional de Investigaciones Científicas y Técnicas
School of Sciences Instituto Nacional de Medicina Tropical
Tezpur University Puerto Iguazú, Argentina
Tezpur, India
Blair W. Perry
Nadia Mushtaq Department of Biology
Department of Biotechnology University of Texas at Arlington
University of Science and Technology Arlington, Texas
Bannu, Pakistan
Daniel Carvalho Pimenta
Edgar Neri-Castro Laboratório de Bioquímica e Biofísica
Instituto de Biotecnología Instituto Butantan
Universidad Nacional Autónoma de México São Paulo, Brazil
Cuernavaca, México
Adriana S. Pontes
Zachary L. Nikolakis Laboratório de Imunologia Aplicada à Saúde Fundação
Department of Biology Oswaldo Cruz
University of Texas at Arlington Unidade Rondônia
Arlington, Texas Porto Velho, Brazil

George Oosthuizen Lucas C. Porta

Department of Surgery Department of Pharmacology
Ngwelezana Hospital Escola Paulista de Medicina
University of Kwa-Zulu Natal, South Africa Universidade Federal de São Paulo
São Paulo, Brazil
Richard W. Orton
Department of Biology Avi Priel
University of Texas at Arlington The Institute for Drug Research
Arlington, Texas School of Pharmacy
The Hebrew University of Jerusalem
Mauro V. Paloschi Jerusalem, Israel
Laboratório de Imunologia Aplicada à Saúde Fundação
Oswaldo Cruz Dileep Punde
Unidade Rondônia Punde Hospital
Porto Velho, Brazil Ashoknagar, India
Contributors xvii

Emelyn Salazar Ene Siigur

National Natural Toxins Research Center National Institute of Chemical Physics and Biophysics
Texas A&M University-Kingsville Tallinn, Estonia
Kingsville, Texas
Jüri Siigur
Andrés Sánchez National Institute of Chemical Physics and Biophysics
Instituto Clodomiro Picado Tallinn, Estonia
Universidad de Costa Rica
San José, Costa Rica Vinícius Lucatelle da Silva
Structural Biology Group
Elda E. Sánchez Multiuser Center for Biomolecular Innovation
National Natural Toxins Research Center Department of Physics, IBILCE/UNESP
Texas A&M University-Kingsville São José do Rio Preto, Brazil
Kingsville, Texas
Cara F. Smith
Laura Sánchez
School of Biological Sciences
Instituto Clodomiro Picado
University of Northern Colorado
Universidad de Costa Rica
Greeley, Colorado
San José, Costa Rica

Melvin Sánchez Andreimar M. Soares

Instituto Clodomiro Picado Centro de Estudos de Biomoléculas Aplicadas à Saúde
Universidad de Costa Rica Fundação Oswaldo Cruz
San José, Costa Rica Unidade Rondônia
and
Marcelo Larami Santoro Departamento de Medicina
Instituto Butantan Universidade Federal de Rondônia
São Paulo, Brazil and
Centro Universitário São Lucas
Porto Velho, Brazil
Anthony J. Saviola
Department of Biochemistry and Molecular Genetics
School of Medicine, University of Colorado Juliana C. Sobrinho
Aurora, Colorado LaBioProt, CEBio, Fundação Oswaldo Cruz
Unidade Rondônia and Departamento de Medicina
Maria Rosa Chitolina Schetinger Universidade Federal de Rondônia
Department of Bioquimica & Toxicologica Porto Velho, Brazil
Federal University Santa Maria
Rio Grande do Sul, Brazil Patrick Jack Spencer
Centro de Biotecnologia
Drew R. Schield Instituto de Pesquisas Energéticas e Nucleares
Department of Biology São Paulo, Brazil
University of Texas at Arlington
Arlington, Texas
Samantha Stack
Department of Medical Microbiology and Immunology, and
Álvaro Segura
Norris Comprehensive Cancer Center
Instituto Clodomiro Picado
Keck School of Medicine
Universidad de Costa Rica
University of Southern California
San José, Costa Rica
Los Angeles, California
Sulamita S. Setubal
Laboratório de Imunologia Aplicada à Saúde Fundação Montamas Suntravat
Oswaldo Cruz National Natural Toxins Research Center
Unidade Rondônia Texas A&M University-Kingsville
Porto Velho, Brazil Kingsville, Texas
xviii Contributors

Stephen D. Swenson Scott A. Weinstein

Department of Neurological Surgery and Department of Department of Toxinology
Biochemistry and Molecular Biology Women’s and Children’s Hospital
Keck School of Medicine Adelaide, Australia
University of Southern California
Los Angeles, California Aundrea K. Westfall
Department of Biology
Zoltan Takacs University of Texas at Arlington
ToxinTech, LLC Arlington, Texas
New York, New York
Kristen L. Wiley
Choo Hock Tan Kentucky Reptile Zoo
Department of Pharmacology Slade, Kentucky
Faculty of Medicine
University of Malaya
Darryl Wood
Kuala Lumpur, Malaysia
Emergency Medicine
Queens Hospital, London
Kae Yi Tan
Queen Mary University London, UK
Department of Molecular Medicine
University of Malaya
Kuala Lumpur, Malaysia Wolfgang Wüster
Molecular Ecology and Fisheries Genetics
Nget Hong Tan Laboratory
Department of Molecular Medicine School of Natural Sciences
University of Malaya Bangor University
Kuala Lumpur, Malaysia Wales, UK

Colin R. Tilbury Michelle Khai Khun Yap

Department of Botany and Zoology School of Science
University of Stellenbosch Monash University Malaysia
Western Cape, South Africa Selangor, Malaysia

Francisco Torres Fernando B. Zanchi

National Natural Toxins Research Center Centro de Estudos de Biomoléculas Aplicadas à Saúde
Texas A&M University-Kingsville Fundação Oswaldo Cruz
Kingsville, Texas Unidade Rondônia
and
Mariángela Vargas Departamento de Medicina
Instituto Clodomiro Picado Universidade Federal de Rondônia
Universidad de Costa Rica Porto Velho, Brazil
San José, Costa Rica
Juliana P. Zuliani
Mauren Villalta Laboratório de Imunologia Aplicada à Saúde Fundação
Instituto Clodomiro Picado Oswaldo Cruz
Universidad de Costa Rica Unidade Rondônia
San José, Costa Rica Porto Velho, Brazil
and
Alejandro Villar-Briones Centro de Estudos de Biomoléculas Aplicadas à Saúde,
Research Support Division Fundação Oswaldo Cruz
Okinawa Institute of Science and Technology Unidade Rondônia
Okinawa-ken, Japan and
Departamento de Medicina
Carl-Wilhelm Vogel Universidade Federal de Rondônia
University of Hawaii Cancer Center Porto Velho, Brazil
Honolulu, Hawaii
Section I
Introduction and Technologies Used in Toxinology

The nicotinic acetylcholine receptor of Torpedo marmorata (PDB 2BG9). The alpha subunits (gold) of the vertebrate skeletal muscle recep-
tor are a common target of neurotoxic snake venom three-finger toxins (drawn with Biovia Discovery Studio 2017R2).