Venom

Venom or zootoxin is a type of

Venoms are often complex mixtures of toxins of differing types. Toxins from venom are used to treat a wide range of medical conditions including thrombosis, arthritis, and some cancers. Studies in venomics are investigating the potential use of venom toxins for many other conditions.

Evolution

The use of venom across a wide variety of

Venoms cause their biological effects via the many toxins that they contain; some venoms are complex mixtures of toxins of differing types. Major classes of toxin in venoms include:^[7]

• serine proteases.^[8]
• Neurotoxins, which primarily affect the nervous systems of animals, such as ion channel toxins. These are found in many venomous taxa, including black widow spiders, scorpions, box jellyfish, cone snails, centipedes and blue-ringed octopuses.^[9]
• Myotoxins, which damage muscles by binding to a receptor. These small, basic peptides are found in snake (such as rattlesnake) and lizard venoms.^{[10][11][12][13]}
• Cytotoxins, which kill individual cells and are found in the apitoxin of honey bees and the venom of black widow spiders.^[14][15]

Taxonomic range

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 on their chelicerae to inject venom, and centipedes, which use forcipules — modified legs — to deliver venom, while scorpions and stinging insects inject venom with a sting. In bees and wasps, the stinger is a modified ovipositor (egg-laying device). In Polistes fuscatus, the female continuously releases a venom that contains a sex pheromone that induces copulatory behavior in males.^[16] In wasps such as Polistes exclamans, venom is used as an alarm pheromone, coordinating a response from the nest and attracting nearby wasps to attack the predator.^[17] In some species, such as Parischnogaster striatula, venom is applied all over the body as an antimicrobial protection.^[18]

Many

There are venomous invertebrates in several

Venom is found in some 200 cartilaginous fishes, including

Some salamanders can extrude sharp venom-tipped ribs.^[28][29] Two frog species in Brazil have tiny spines around the crown of their skulls which, on impact, deliver venom into their targets.^[30]

Reptiles

The venom of the prairie rattlesnake, Crotalus viridis (left), includes metalloproteinases (example on the right) which help digest prey before eating.

Some 450 species of snake are venomous.

Venom is found in a few other reptiles such as the Mexican beaded lizard,^[35] the gila monster,^[36] and some monitor lizards, including the Komodo dragon.^[37] Mass spectrometry showed that the mixture of proteins present in their venom is as complex as the mixture of proteins found in snake venom.^[37][38] Some lizards possess a venom gland; they form a hypothetical clade,

Euchambersia, an extinct genus of therocephalians, is hypothesized to have had venom glands attached to its canine teeth.^[40]

A few species of living mammals are venomous, including

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 was once thought.[44] 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 of why venom has become a homoplastic trait and why very different animals have convergently evolved.^[13]

Venom and humans

Envenomation resulted in 57,000 human deaths in 2013, down from 76,000 deaths in 1990.^[45] Venoms, found in over 173,000 species, have potential to treat a wide range of diseases, explored in over 5,000 scientific papers.^[36]

In medicine, snake venom proteins are used to treat conditions including thrombosis, arthritis, and some cancers.^[46][47] Gila monster venom contains exenatide, used to treat type 2 diabetes.^[36] Solenopsins extracted from fire ant venom has demonstrated biomedical applications, ranging from cancer treatment to psoriasis.^[48][49] A branch of science, venomics, has been established to study the proteins associated with venom and how individual components of venom can be used for pharmaceutical means.^[50]

Resistance

Northern Pacific rattlesnake

's powerful venom.

Venom is used as a trophic weapon by many predator species. The coevolution between predators and prey is the driving force of venom resistance, which has evolved multiple times throughout the animal kingdom.^[51] The coevolution between venomous predators and venom-resistant prey has been described as a chemical arms race.^[52] Predator/prey pairs are expected to coevolve over long periods of time.^[53] As the predator capitalizes on susceptible individuals, the surviving individuals are limited to those able to evade predation.^[54] Resistance typically increases over time as the predator becomes increasingly unable to subdue resistant prey.^[55] The cost of developing venom resistance is high for both predator and prey.^[56] The payoff for the cost of physiological resistance is an increased chance of survival for prey, but it allows predators to expand into underutilised trophic niches.^[57]

The

Northern Pacific rattlesnake.^[58] The resistance involves toxin scavenging and depends on the population. Where rattlesnake populations are denser, squirrel resistance is higher.^[59] Rattlesnakes have responded locally by increasing the effectiveness of their venom.^[60]

The kingsnakes of the Americas are constrictors that prey on many venomous snakes.^[61] They have evolved resistance which does not vary with age or exposure.^[55] They are immune to the venom of snakes in their immediate environment, like copperheads, cottonmouths, and North American rattlesnakes, but not to the venom of, for example, king cobras or black mambas.^[62]

Among marine animals, eels are resistant to sea snake venoms, which contain complex mixtures of neurotoxins, myotoxins, and nephrotoxins, varying according to species.^[63][64] Eels are especially resistant to the venom of sea snakes that specialise in feeding on them, implying coevolution; non-prey fishes have little resistance to sea snake venom.^[65]

Clownfish always live among the tentacles of venomous

nematocysts and mucous secretions. The toxins are composed of peptides and proteins. They are used to acquire prey and to deter predators by causing pain, loss of muscular coordination, and tissue damage. Clownfish have a protective mucus that acts as a chemical camouflage or macromolecular mimicry preventing "not self" recognition by the sea anemone and nematocyst discharge.^[71][72][73] Clownfish may acclimate their mucus to resemble that of a specific species of sea anemone.^[73]

See also

• Schmidt Sting Pain Index

References

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