Venom

Wasp sting, with a droplet of venom

Venom is a secretion containing one or more toxins produced by an animal to cause harm to another.[1] 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; myotoxins, which damage muscles.

Venomous animals kill tens of thousands of humans per year. However, the toxins in many venoms have potential to treat a wide range of diseases.

Evolution

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.[2]

Mechanisms

Phospholipase A2, an enzyme found in bee stings.

Venoms 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:[3]

Taxonomic range

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.[12] 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.[13] In Dolichovespula arenaria, the observed spraying of venom out of their sting has been seen from workers in large colonies.[14] In other cases like Parischnogaster striatula, the venom is applied all over their body as an antimicrobial protection.[15] The venom from Agelaia pallipes has inhibitory effects on processes like chemotaxis and hemolysis which can lead to organ failure.[16]

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.[17]

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. At least one ant species (Polyrhachis dives) has been shown to use venom topically for the sterilisation of pathogens.[18]

Other invertebrates

There are venomous invertebrates in several phyla, including jellyfish such as the dangerous box jellyfish[19] and sea anemones among the Cnidaria,[20] sea urchins among the Echinodermata,[21] and cone snails[22] and cephalopods including octopuses among the Molluscs.[23]

Vertebrates

Fish

Venom is found in some 200 cartilaginous fishes – 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.[24]

Amphibians

Among amphibians, some salamanders can extrude sharp venom-tipped ribs.[25][26] The poison dart frogs (Dendrobatidae) secret toxic alkaloids including batrachotoxins, histrionicotoxins, indolizidines, pumiliotoxins, decahydroquinolines, gephyrotoxins, 2,6-disubstituted piperidines, 2,5-disubstituted pyrrolidines, pyridyl-piperidines, indole alkaloids, azatricyclododecenes and amidine alkaloids from the granular glands in their skin.[27]

Reptiles

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.[24] 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.[28] 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.[29] Snake venom may have originated with duplication of genes that had been expressed in the salivary glands of ancestors.[30][31]

Venom is found in a few other reptiles such as the Mexican beaded lizard,[32] the gila monster,[33] and some monitor lizards including the Komodo dragon.[34] Mass spectrometry showed that the mixture of proteins present in the venom is as complex as the proteins found in snake venom.[34][35] 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.[36]

Mammals

Euchambersia, an extinct genus of therocephalians, is hypothesized to have had venom glands attached to its canine teeth.[37]

A few species of living mammals are venomous, including solenodons, shrews, vampire bats, the male platypus and the slow loris.[24][38] Shrews are known to have venomous saliva and most likely evolved their trait similarly to snakes.[39] 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.[40]

Extensive 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.[41] 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.[42]

Venom and humans

Venomous animals resulted in 57,000 human deaths in 2013, down from 76,000 deaths in 1990.[43]

Venoms, found in over 173,000 species, have potential to treat a wide range of diseases, explored in over 5,000 scientific papers.[33] Snake venoms contain proteins which can be used to treat conditions including thrombosis, arthritis, and some cancers.[44][45] Gila monster venom contains exenatide, used to treat type 2 diabetes.[33]

See also

References

  1. ↑ "venom" at Dorland's Medical Dictionary
  2. ↑ Kordiš D.; Gubenšek F. (2000). "Adaptive evolution of animal toxin multigene families". Gene. 261 (1): 43–52. doi:10.1016/s0378-1119(00)00490-x. PMID 11164036.
  3. ↑ Harris, J. B. (September 2004). "Animal poisons and the nervous system: what the neurologist needs to know". Journal of Neurology, Neurosurgery & Psychiatry. 75 (suppl_3): iii40–iii46. doi:10.1136/jnnp.2004.045724. PMC 1765666. PMID 15316044.
  4. ↑ Raffray M, Cohen GM; Cohen (1997). "Apoptosis and necrosis in toxicology: a continuum or distinct modes of cell death?". Pharmacol. Ther. 75 (3): 153–77. doi:10.1016/s0163-7258(97)00037-5. PMID 9504137.
  5. ↑ Dutertre, Sébastien; Lewis, Richard J. (2006). "Toxin insights into nicotinic acetylcholine receptors". Biochemical Pharmacology. 72 (6): 661–670. doi:10.1016/j.bcp.2006.03.027. PMID 16716265.
  6. ↑ Nicastro, G; Franzoni, L; de Chiara, C; Mancin, A. C.; Giglio, J. R.; Spisni, A. (May 2003). "Solution structure of crotamine, a Na+ channel affecting toxin from Crotalus durissus terrificus venom". Eur. J. Biochem. 270 (9): 1969–79. doi:10.1046/j.1432-1033.2003.03563.x. PMID 12709056.
  7. ↑ Griffin P. R.; Aird S. D. (1990). "A new small myotoxin from the venom of the prairie rattlesnake (Crotalus viridis viridis)". FEBS Lett. 274 (1): 43–47. doi:10.1016/0014-5793(90)81325-I. PMID 2253781.
  8. ↑ Samejima Y.; Aoki, Y; Mebs, D. (1991). "Amino acid sequence of a myotoxin from venom of the eastern diamondback rattlesnake (Crotalus adamanteus)". Toxicon. 29 (4): 461–468. doi:10.1016/0041-0101(91)90020-r. PMID 1862521.
  9. ↑ 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. (June 2008). "Defensins and the convergent evolution of platypus and reptile venom genes". Genome Research. 18 (6): 986–94. doi:10.1101/gr.7149808. PMC 2413166. PMID 18463304.
  10. ↑ Sobral, Filipa; Sampaio, Andreia; Falcão, Soraia; Queiroz, Maria João R.P.; Calhelha, Ricardo C.; Vilas-Boas, Miguel; Ferreira, Isabel C.F.R. (2016). "Chemical characterization, antioxidant, anti-inflammatory and cytotoxic properties of bee venom collected in Northeast Portugal". Food and Chemical Toxicology. 94: 172–177. doi:10.1016/j.fct.2016.06.008. hdl:10198/13492. PMID 27288930.
  11. ↑ Peng, Xiaozhen; Dai, Zhipan; Lei, Qian; Liang, Long; Yan, Shuai; Wang, Xianchun (April 2017). "Cytotoxic and apoptotic activities of black widow spiderling extract against HeLa cells". Experimental and Therapeutic Medicine. 13 (6): 3267–3274. doi:10.3892/etm.2017.4391. PMC 5450530. PMID 28587399.
  12. ↑ 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. doi:10.1007/bf00988043. PMID 24407344.
  13. ↑ Post Downing, Jeanne (1984). "Alarm response to venom by social wasps Polistes exclamans and P. fuscatus". Journal of Chemical Ecology. 10 (10): 1425–1433. doi:10.1007/BF00990313. PMID 24318343.
  14. ↑ Greene, Alex. "The Aerial Yellowjacket Dolichovespula Arenaria." Academia.edu. Department of Entomology — Washington State University, n.d. Web. 25 Sept. 2014.
  15. ↑ Baracchi, David (January 2012). "From individual to collective immunity: The role of the venom as antimicrobial agent in the Stenogastrinae wasp societies". Journal of Insect Physiology. 58 (1): 188–193. doi:10.1016/j.jinsphys.2011.11.007. hdl:2158/790328. PMID 22108024.
  16. ↑ Baptista-Saidemberg Nicoli; et al. (2011). "Profiling the peptidome of the venom from the social wasp Agelaia pallipes pallipes". Journal of Proteomics. 74 (10): 2123–2137. doi:10.1016/j.jprot.2011.06.004. PMID 21693203.
  17. ↑ Pinto, Antônio F.M.; Berger, Markus; Reck, José; Terra, Renata M.S.; Guimarães, Jorge A. (15 December 2010). "Lonomia obliqua venom: In vivo effects and molecular aspects associated with the hemorrhagic syndrome". Toxicon : Official Journal of the International Society on Toxinology. 56 (7): 1103–1112. doi:10.1016/j.toxicon.2010.01.013. PMID 20114060.
  18. ↑ Graystock, Peter; Hughes, William O. H. (2011). "Disease resistance in a weaver ant, Polyrhachis dives, and the role of antibiotic-producing glands". Behavioral Ecology and Sociobiology. 65 (12): 2319–2327. doi:10.1007/s00265-011-1242-y.
  19. ↑ Frost, Emily (30 August 2013). "What's Behind That Jellyfish Sting?". Smithsonian. Retrieved 30 September 2018.
  20. ↑ Bonamonte, Domenico; Angelini, Gianni (2016). Aquatic Dermatology: Biotic, Chemical and Physical Agents. Springer International. pp. 54–56. ISBN 978-3-319-40615-2.
  21. ↑ Gallagher, Scott A. (2017-08-02). "Echinoderm Envenomation". eMedicine. Retrieved 12 October 2010.
  22. ↑ Olivera, B. M.; Teichert, R. W. (2007). "Diversity of the neurotoxic Conus peptides: a model for concerted pharmacological discovery". Molecular Interventions. 7 (5): 251–260. doi:10.1124/mi.7.5.7. PMID 17932414.
  23. ↑ Barry, Carolyn (17 April 2009). "All Octopuses Are Venomous, Study Says". National Geographic. Retrieved 30 September 2018.
  24. 1 2 3 Smith, William Leo; Wheeler, Ward C. (2006). "Venom Evolution Widespread in Fishes: A Phylogenetic Road Map for the Bioprospecting of Piscine Venoms". Journal of Heredity. 97 (3): 206–217. doi:10.1093/jhered/esj034. PMID 16740627.
  25. ↑ Venomous Amphibians (Page 1) - Reptiles (Including Dinosaurs) and Amphibians - Ask a Biologist Q&A. Askabiologist.org.uk. Retrieved on 2013-07-17.
  26. ↑ Nowak, R. T.; Brodie, E. D. (1978). "Rib Penetration and Associated Antipredator Adaptations in the Salamander Pleurodeles waltl (Salamandridae)". Copeia. 1978 (3): 424–429. doi:10.2307/1443606. JSTOR 1443606.
  27. ↑ Daly, John W.; Myers, Charles W.; Whittaker, Noel (1987). "Further classification of skin alkaloids from neotropical poison frogs (dendrobatidae), with a general survey of toxic/noxious substances in the amphibia". Toxicon. 25 (10): 1023–1095. doi:10.1016/0041-0101(87)90265-0.
  28. ↑ Bauchot, Roland (1994). Snakes: A Natural History. Sterling. pp. 194–209. ISBN 978-1-4027-3181-5.
  29. ↑ "Snake Bites". A. D. A. M. Inc. 16 October 2017. Retrieved 30 September 2018.
  30. ↑ Hargreaves, Adam D.; Swain, Martin T.; Hegarty, Matthew J.; Logan, Darren W.; Mulley, John F. (30 July 2014). "Restriction and Recruitment—Gene Duplication and the Origin and Evolution of Snake Venom Toxins". Genome Biology and Evolution. 6 (8): 2088–2095. doi:10.1093/gbe/evu166. PMC 4231632. PMID 25079342.
  31. ↑ Daltry, Jennifer C.; Wuester, Wolfgang; Thorpe, Roger S. (1996). "Diet and snake venom evolution". Nature. 379 (6565): 537–540. doi:10.1038/379537a0. PMID 8596631.
  32. ↑ Cantrell, F. L. (2003). "Envenomation by the Mexican beaded lizard: a case report". Journal of Toxicology. Clinical Toxicology. 41 (3): 241–244. PMID 12807305.
  33. 1 2 3 Mullin, Emily (29 November 2015). "Animal Venom Database Could Be Boon To Drug Development". Forbes. Retrieved 30 September 2018.
  34. 1 2 Fry BG, Wroe S, Teeuwisse W, et al. (June 2009). "A central role for venom in predation by Varanus komodoensis (Komodo Dragon) and the extinct giant Varanus (Megalania) priscus". PNAS. 106 (22): 8969–74. doi:10.1073/pnas.0810883106. PMC 2690028. PMID 19451641.
  35. ↑ Fry, B. G., W. Wuster, S. F. R. Ramjan, T. Jackson, P. Martelli, and R. M. Kini. 2003c. Analysis of Colubroidea snake venoms by liquid chromatography with mass spectrometry: Evolutionary and toxinological implications. Rapid Communications in Mass Spectrometry 17:2047-2062.
  36. ↑ Fry, B. G.; Vidal, N.; Norman, J. A.; Vonk, F. J.; Scheib, H.; Ramjan, S. F.; Kuruppu, S.; Fung, K.; Hedges, S. B.; Richardson MK; Hodgson WC; Ignjatovic V; Summerhayes R; Kochva, E. (February 2006). "Early evolution of the venom system in lizards and snakes". Nature. 439 (7076): 584–588. doi:10.1038/nature04328. PMID 16292255.
  37. ↑ Benoit, J.; Norton, L.A.; Manger, P.R.; Rubidge, B.S. (2017). "Reappraisal of the envenoming capacity of Euchambersia mirabilis (Therapsida, Therocephalia) using μCT-scanning techniques". PLoS ONE. 12 (2): e0172047. doi:10.1371/journal.pone.0172047. PMC 5302418. PMID 28187210.
  38. ↑ 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. doi:10.1186/1678-9199-19-21. PMC 3852360. PMID 24074353.
  39. ↑ Ligabue-Braun R.; Verli H.; Carlini C. R. (2012). "Venomous mammals: a review". Toxicon. 59 (7–8): 680–695. doi:10.1016/j.toxicon.2012.02.012. PMID 22410495.
  40. ↑ Jørn H. Hurum, Zhe-Xi Luo, and Zofia Kielan-Jaworowska, Were mammals originally venomous?, Acta Palaeontologica Polonica 51 (1), 2006: 1-11
  41. ↑ Wong E. S.; Belov K. (2012). "Venom evolution through gene duplications". Gene. 496 (1): 1–7. doi:10.1016/j.gene.2012.01.009. PMID 22285376.
  42. ↑ 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. doi:10.1101/gr.7149808. PMC 2413166. PMID 18463304.
  43. ↑ GBD 2013 Mortality and Causes of Death, Collaborators (17 December 2014). "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. 385 (9963): 117–71. doi:10.1016/S0140-6736(14)61682-2. PMC 4340604. PMID 25530442.
  44. ↑ Pal, S. K.; Gomes, A.; Dasgupta, S. C.; Gomes, A. (2002). "Snake venom as therapeutic agents: from toxin to drug development". Indian Journal of Experimental Biology. 40 (12): 1353–8. ISSN 0019-5189. PMID 12974396.
  45. ↑ Holland, Jennifer S. (February 2013). "The Bite That Heals". National Geographic. Retrieved 30 September 2018.
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