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venom
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{{About|the class of biotoxins}}{{Use dmy dates|date=October 2013}}File:Waspstinger1658-2.jpg|thumb|upright=1.35|Wasp sting, with a droplet of venom]]Venom is a secretion containing one or more toxins produced by an animal.{{DorlandsDict|eight/000115553|venom}} Venom has evolved in a wide variety of animals, both predators and prey, and both vertebrates and invertebrates.Venoms kill through the action of at least four major classes of toxin, namely necrotoxins and cytotoxins, which kill cells; neurotoxins, which affect nervous systems; and myotoxins, which damage muscles. Biologically, venom is distinguished from poison in that poisons are ingested, while venom is delivered in a bite, sting, or similar action. Venomous animals cause tens of thousands of human deaths per year. However, the toxins in many venoms have potential to treat a wide range of diseases.

Evolution

{{further|Evolution of snake venom}}The use of venom across a wide variety of taxa is an example of convergent evolution. It is difficult to conclude exactly how this trait came to be so intensely widespread and diversified. The multigene families that encode the toxins of venomous animals are actively selected, creating more diverse toxins with specific functions. Venoms adapt to their environment and victims and accordingly evolve to become maximally efficient on a predator's particular prey (particularly the precise ion channels within the prey). Consequently, venoms become specialized to an animal's standard diet.JOURNAL, Kordiš, D., Gubenšek, F., 2000, Adaptive evolution of animal toxin multigene families, Gene, 261, 1, 43–52, 10.1016/s0378-1119(00)00490-x, 11164036,

Mechanisms

File:1poc.png|thumb|right|Phospholipase A2, an enzyme found in bee stingbee stingVenoms cause their biological effects via the toxins that they contain; some venoms are complex mixtures of toxins of differing types. Among the major classes of toxin in venoms are:JOURNAL, Harris, J. B., Animal poisons and the nervous system: what the neurologist needs to know, Journal of Neurology, Neurosurgery & Psychiatry, 75, suppl_3, September 2004, 10.1136/jnnp.2004.045724, 15316044, 1765666, iii40–iii46,
  • Necrotoxins, which cause necrosis (i.e., death) in the cells they encounter.JOURNAL, Raffray M, Cohen GM, Apoptosis and necrosis in toxicology: a continuum or distinct modes of cell death?, Pharmacol. Ther., 75, 3, 153–177, 1997, 9504137, 10.1016/s0163-7258(97)00037-5, Cohen, The venom of most viper species contains phospholipase and trypsin-like serine proteases.
  • Neurotoxins, which primarily affect the nervous systems of animals.JOURNAL, Dutertre, Sébastien, Lewis, Richard J., Toxin insights into nicotinic acetylcholine receptors, Biochemical Pharmacology, 72, 6, 2006, 10.1016/j.bcp.2006.03.027, 16716265, 661–670, These include (:Category:Ion channel toxins|ion channel toxins) that disrupt ion channel conductance. Black widow spider, scorpion, box jellyfish, cone snail, centipede and blue-ringed octopus venoms (among many others) function in this way.
  • Myotoxins, which damage muscles by binding to a receptor, are small, basic peptides found in snake (such as rattlesnake) and lizard venoms.JOURNAL, Nicastro, G; Franzoni, L; de Chiara, C; Mancin, A. C.; Giglio, J. R.; Spisni, A., Solution structure of crotamine, a Na+ channel affecting toxin from Crotalus durissus terrificus venom, Eur. J. Biochem., 270, 9, 1969–1979, May 2003, 12709056, 10.1046/j.1432-1033.2003.03563.x, JOURNAL, Griffin, P. R.; Aird, S. D., A new small myotoxin from the venom of the prairie rattlesnake (Crotalus viridis viridis), FEBS Lett., 274, 1, 43–47, 1990, 2253781, 10.1016/0014-5793(90)81325-I, JOURNAL, Samejima Y.; Aoki, Y; Mebs, D., Amino acid sequence of a myotoxin from venom of the eastern diamondback rattlesnake (Crotalus adamanteus), Toxicon, 29, 4, 461–468, 1991, 1862521, 10.1016/0041-0101(91)90020-r, JOURNAL, Whittington, C. M., Papenfuss, A. T., Bansal, P., Torres, A. M., Wong, E. S., Deakin, J. E., Graves, T., Alsop, A., Schatzkamer, K., Kremitzki, C., Ponting, C. P., Temple-Smith, P., Warren, W. C., Kuchel, P. W., Belov, K., Defensins and the convergent evolution of platypus and reptile venom genes, Genome Research, June 2008, 18, 6, 986–094, 18463304, 10.1101/gr.7149808, 2413166,
  • Cytotoxins, which kill individual cells, are found in the apitoxin of honey bees and the venom of black widow spiders.JOURNAL, Sobral, Filipa, Sampaio, Andreia, Falcão, Soraia, Queiroz, Maria João R.P., Calhelha, Ricardo C., Vilas-Boas, Miguel, Ferreira, Isabel C.F.R., Chemical characterization, antioxidant, anti-inflammatory and cytotoxic properties of bee venom collected in Northeast Portugal, Food and Chemical Toxicology, 94, 2016, 10.1016/j.fct.2016.06.008, 27288930, 172–177, 10198/13492,weblink JOURNAL, Peng, Xiaozhen, Dai, Zhipan, Lei, Qian, Liang, Long, Yan, Shuai, Wang, Xianchun, Cytotoxic and apoptotic activities of black widow spiderling extract against HeLa cells, Experimental and Therapeutic Medicine, 13, 6, April 2017, 10.3892/etm.2017.4391, 28587399, 5450530, 3267–3274,

Taxonomic range

(file:Venomous Animals.jpg|thumb|Venomous animals)Venom is widely distributed taxonomically, being found in both invertebrates and vertebrates; in aquatic and terrestrial animals; and among both predators and prey. The major groups of venomous animals are described below.

Arthropods

Venomous arthropods include spiders, which use fangs — part of their chelicerae — to inject venom; and centipedes, which use forcipules — modified legs — to deliver venom; along with scorpions and stinging insects, which inject venom with a sting.In insects such as bees and wasps, the stinger is a modified egg-laying device — the ovipositor. In Polistes fuscatus, the female continuously releases a venom that contains a sex pheromone that induces copulatory behavior in males.JOURNAL, Post David, Jeanne Robert, 1983, Venom: Source of a Sex Pheromone in the Social Wasp Polistes fuscatus (Hymenoptera: Vespidae), Journal of Chemical Ecology, 9, 2, 259–266, 10.1007/bf00988043, 24407344, In Polistes exclamans, venom is used as an alarm pheromone, coordinating a response with from the nest and attracting nearby wasps to attack the predator.JOURNAL, Post Downing, Jeanne, 1984, Alarm response to venom by social wasps Polistes exclamans and P. fuscatus, Journal of Chemical Ecology, 10, 10, 1425–1433, 10.1007/BF00990313, 24318343, In Dolichovespula arenaria, the observed spraying of venom out of their sting has been seen from workers in large colonies.Greene, Alex. "The Aerial Yellowjacket Dolichovespula Arenaria." Academia.edu. Department of Entomology — Washington State University, n.d. Web. 25 September 2014. In other cases like Parischnogaster striatula, the venom is applied all over their body as an antimicrobial protection.JOURNAL, From individual to collective immunity: The role of the venom as antimicrobial agent in the Stenogastrinae wasp societies, Baracchi, David, January 2012, Journal of Insect Physiology, 10.1016/j.jinsphys.2011.11.007, 22108024, 58, 1, 188–193, 2158/790328, The venom from Agelaia pallipes has inhibitory effects on processes like chemotaxis and hemolysis which can lead to organ failure.JOURNAL, Baptista-Saidemberg, Nicoli, 2011, Profiling the peptidome of the venom from the social wasp Agelaia pallipes pallipes, Journal of Proteomics, 74, 10, 2123–2137, 10.1016/j.jprot.2011.06.004, 21693203, etal, Many caterpillars have defensive venom glands associated with specialized bristles on the body, known as urticating hairs, which can be lethal to humans (e.g., that of the Lonomia moth), although the venom's strength varies depending on the species.JOURNAL, Pinto, Antônio F. M., Berger, Markus, Reck, José, Terra, Renata M. S., Guimarães, Jorge A., Lonomia obliqua venom: In vivo effects and molecular aspects associated with the hemorrhagic syndrome, Toxicon, 56, 7, 15 December 2010, 20114060, 10.1016/j.toxicon.2010.01.013, 1103–1112, Bees synthesize and employ an acidic venom (apitoxin) to cause pain in those that they sting to defend their hives and food stores, whereas wasps use a chemically different alkaline venom designed to paralyze prey, so it can be stored alive in the food chambers of their young. The use of venom is much more widespread than just these examples. Other insects, such as true bugs and many ants, also produce venom.JOURNAL, Touchard, Axel, Aili, Samira, Fox, Eduardo, Escoubas, Pierre, Orivel, Jérôme, Nicholson, Graham, Dejean, Alain, 2016-01-20, The Biochemical Toxin Arsenal from Ant Venoms, Toxins, 8, 1, 30, 10.3390/toxins8010030, 26805882, 4728552, 2072-6651, At least one ant species (Polyrhachis dives) has been shown to use venom topically for the sterilisation of pathogens.JOURNAL, Graystock, Peter, Hughes, William O. H., Disease resistance in a weaver ant, Polyrhachis dives, and the role of antibiotic-producing glands, Behavioral Ecology and Sociobiology, 2011, 10.1007/s00265-011-1242-y, 65, 12, 2319–2327,

Other invertebrates

There are venomous invertebrates in several phyla, including jellyfish such as the dangerous box jellyfishMAGAZINE, Frost, Emily, What's Behind That Jellyfish Sting?,weblink Smithsonian, 30 September 2018, 30 August 2013, and sea anemones among the Cnidaria,BOOK, Bonamonte, Domenico, Angelini, Gianni, Aquatic Dermatology: Biotic, Chemical and Physical Agents,weblink 2016, Springer International, 978-3-319-40615-2, 54–56, sea urchins among the Echinodermata,JOURNAL, Gallagher, Scott A., Echinoderm Envenomation,weblink EMedicine, 12 October 2010, 2017-08-02, and cone snailsJOURNAL, Olivera, B. M., Teichert, R. W., Diversity of the neurotoxic Conus peptides: a model for concerted pharmacological discovery, Molecular Interventions, 2007, 7, 5, 251–260, 17932414, 10.1124/mi.7.5.7, and cephalopods including octopuses among the Molluscs.MAGAZINE, Barry, Carolyn, All Octopuses Are Venomous, Study Says,weblink National Geographic, 30 September 2018, 17 April 2009,

Vertebrates

Fish

Venom is found in some 200 cartilaginous fishes, including stingrays, sharks, and chimaeras; the catfishes (about 1000 venomous species); and 11 clades of spiny-rayed fishes (Acanthomorpha), containing the scorpionfishes (over 300 species), stonefishes (over 80 species), gurnard perches, blennies, rabbitfishes, surgeonfishes, some velvetfishes, some toadfishes, coral crouchers, red velvetfishes, scats, rockfishes, deepwater scorpionfishes, waspfishes, weevers, and stargazers.JOURNAL, Smith, William Leo, Wheeler, Ward C., Venom Evolution Widespread in Fishes: A Phylogenetic Road Map for the Bioprospecting of Piscine Venoms, Journal of Heredity, 97, 3, 2006, 10.1093/jhered/esj034, 16740627, 206–217,

Amphibians

Among amphibians, some salamanders can extrude sharp venom-tipped ribs.Venomous Amphibians (Page 1) - Reptiles (Including Dinosaurs) and Amphibians - Ask a Biologist Q&A. Askabiologist.org.uk. Retrieved on 2013-07-17.JOURNAL, Nowak, R. T., Brodie, E. D., Rib Penetration and Associated Antipredator Adaptations in the Salamander Pleurodeles waltl (Salamandridae), Copeia, 1978, 3, 424–429, 1978, 10.2307/1443606, 1443606,

Reptiles

{{multiple image| align =right| width1 =160|image1=PrairieRattlesnake.jpg| width2=144|image2=PDB 1rm8 EBI.jpg|footer=The venom of the prairie rattlesnake, Crotalus viridis (left) includes metalloproteinases (example on the right) which help digest the prey before the snake eats it. }}Some 450 species of snake are venomous. Snake venom is produced by glands below the eye (the mandibular gland) and delivered to the victim through tubular or channeled fangs. Snake venoms contain a variety of peptide toxins, including proteases, which hydrolyze protein peptide bonds, nucleases, which hydrolyze the phosphodiester bonds of DNA, and neurotoxins, which disable signalling in the nervous system.BOOK, Bauchot, Roland, Snakes: A Natural History, 1994, Sterling, 978-1-4027-3181-5, 194–209, Snake venom causes symptoms including pain, swelling, tissue necrosis, low blood pressure, convulsions, hemorrhage (varying by species of snake), respiratory paralysis, kidney failure, coma and death.WEB, Snake Bites,weblink A. D. A. M. Inc, 30 September 2018, 16 October 2017, Snake venom may have originated with duplication of genes that had been expressed in the salivary glands of ancestors.JOURNAL, Hargreaves, Adam D., Swain, Martin T., Hegarty, Matthew J., Logan, Darren W., Mulley, John F., Restriction and Recruitment—Gene Duplication and the Origin and Evolution of Snake Venom Toxins, Genome Biology and Evolution, 6, 8, 30 July 2014, 10.1093/gbe/evu166, 25079342, 4231632, 2088–2095, JOURNAL, Daltry, Jennifer C., Wuester, Wolfgang, Thorpe, Roger S., 1996, Diet and snake venom evolution, Nature, 379, 6565, 537–540, 10.1038/379537a0, 8596631, Venom is found in a few other reptiles such as the Mexican beaded lizard,JOURNAL, Cantrell, F. L., Envenomation by the Mexican beaded lizard: a case report, Journal of Toxicology. Clinical Toxicology, 41, 3, 2003, 12807305, 241–244, 10.1081/CLT-120021105, the gila monster, and some monitor lizards including the Komodo dragon.JOURNAL, Fry, B. G., Wroe, S., Teeuwisse, W., etal, A central role for venom in predation by Varanus komodoensis (Komodo Dragon) and the extinct giant Varanus (Megalania) priscus, PNAS, 106, 22, 8969–8974, June 2009, 19451641, 2690028, 10.1073/pnas.0810883106, Mass spectrometry showed that the mixture of proteins present in their venom is as complex as the mixture of proteins found in snake venom.Fry, B. G.; Wuster, W.; Ramjan, S. F. R.; Jackson, T.; Martelli, P.; Kini, R. M. 2003c. Analysis of Colubroidea snake venoms by liquid chromatography with mass spectrometry: Evolutionary and toxinological implications. Rapid Communications in Mass Spectrometry 17:2047-2062.Some lizards possess a venom gland; they form a hypothetical clade, Toxicofera, containing the suborders Serpentes and Iguania and the families Varanidae, Anguidae, and Helodermatidae.JOURNAL, February 2006, Early evolution of the venom system in lizards and snakes, Nature, 439, 584–588, 10.1038/nature04328, 16292255, Fry, B. G., Vidal, N., Norman, J. A., Vonk, F. J., Scheib, H., Ramjan, S. F., Kuruppu, S., Fung, K., Hedges, S. B., Richardson, M. K., Hodgson, W. C., Ignjatovic, V., Summerhayes, R., Kochva, E., 7076,

Mammals

Euchambersia, an extinct genus of therocephalians, is hypothesized to have had venom glands attached to its canine teeth.JOURNAL, Benoit, J., Norton, L. A., Manger, P. R., Rubidge, B. S., Reappraisal of the envenoming capacity of Euchambersia mirabilis (Therapsida, Therocephalia) using μCT-scanning techniques, 2017, PLoS ONE, 12, 2, e0172047, 10.1371/journal.pone.0172047, 28187210, 5302418, A few species of living mammals are venomous, including solenodons, shrews, vampire bats, the male platypus and the slow loris.JOURNAL, Nekaris, K. Anne-Isola, Moore, Richard S., Rode, E. Johanna, Fry, Bryan G., 2013-09-27, Mad, bad and dangerous to know: the biochemistry, ecology and evolution of slow loris venom, Journal of Venomous Animals and Toxins Including Tropical Diseases, 19, 1, 21, 10.1186/1678-9199-19-21, 24074353, 3852360, Shrews are known to have venomous saliva and most likely evolved their trait similarly to snakes.JOURNAL, Ligabue-Braun, R., Verli, H., Carlini, C. R., 2012, Venomous mammals: a review, Toxicon, 59, 7–8, 680–695, 10.1016/j.toxicon.2012.02.012, 22410495, The presence of tarsal spurs akin to those of the platypus in many non-therian Mammaliaformes groups suggests that venom was an ancestral characteristic among mammals.Jørn H. Hurum, Zhe-Xi Luo, and Zofia Kielan-Jaworowska, Were mammals originally venomous?, Acta Palaeontologica Polonica 51 (1), 2006: 1-11Extensive research on platypuses shows that their toxin was initially formed from gene duplication, but data provides evidence that the further evolution of platypus venom does not rely as much on gene duplication as once was thought.JOURNAL, Wong, E. S., Belov, K., 2012, Venom evolution through gene duplications, Gene, 496, 1, 1–7, 10.1016/j.gene.2012.01.009, 22285376, Modified sweat glands are what evolved into platypus venom glands. Although it is proven that reptile and platypus venom have independently evolved, it is thought that there are certain protein structures that are favored to evolve into toxic molecules. This provides more evidence as to why venom has become a homoplastic trait and why very different animals have convergently evolved.JOURNAL, Whittington C. M., Papenfuss A. T., Bansal P., Torres A. M., Wong E. S., Deakin J. E., Belov K., 2008, Defensins and the convergent evolution of platypus and reptile venom genes, Genome Research, 18, 6, 986–994, 10.1101/gr.7149808, 18463304, 2413166,

Venom and humans

Venomous animals resulted in 57,000 human deaths in 2013, down from 76,000 deaths in 1990.JOURNAL, GBD 2013 Mortality and Causes of Death, Collaborators, Global, regional, and national age-sex specific all-cause and cause-specific mortality for 240 causes of death, 1990-2013: a systematic analysis for the Global Burden of Disease Study 2013, Lancet, 17 December 2014, 25530442, 10.1016/S0140-6736(14)61682-2, 4340604, 385, 9963, 117–171, Venoms, found in over 173,000 species, have potential to treat a wide range of diseases, explored in over 5,000 scientific papers.MAGAZINE, Mullin, Emily, Animal Venom Database Could Be Boon To Drug Development,weblink Forbes, 30 September 2018, 29 November 2015, Snake venoms contain proteins which can be used to treat conditions including thrombosis, arthritis, and some cancers.JOURNAL, Pal, S. K., Gomes, A., Dasgupta, S. C., Gomes, A., Snake venom as therapeutic agents: from toxin to drug development., Indian Journal of Experimental Biology, 40, 12, 2002, 12974396, 1353–1358, MAGAZINE, Holland, Jennifer S., The Bite That Heals,weblink National Geographic, 30 September 2018, February 2013, Gila monster venom contains exenatide, used to treat type 2 diabetes.Solenopsins extracted from fire ant venom has demonstrated biomedical applications, ranging from cancer treatment to psoriasis.JOURNAL, Fox, Eduardo G.P., Xu, Meng, Wang, Lei, Chen, Li, Lu, Yong-Yue, May 2018, Speedy milking of fresh venom from aculeate hymenopterans, Toxicon, 146, 120–123, 10.1016/j.toxicon.2018.02.050, 29510162, {{Citation|last=Fox|first=Eduardo Gonçalves Paterson|title=Venom Toxins of Fire Ants|date=2021|work=Venom Genomics and Proteomics: Venom Genomics and Proteomics|pages=1–16|editor-last=Gopalakrishnakone|editor-first=P.|publisher=Springer Netherlands|doi=10.1007/978-94-007-6649-5_38-1|isbn=9789400766495|editor2-last=Calvete|editor2-first=Juan J.|doi-broken-date=2019-08-20}}

See also

References

{{reflist}}{{Toxins}}{{Toxicology}}{{Poisoning and toxicity}}

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