Venom

12/14/2019 Venom - Wikipedia
Venom
Venom is a secretion containing one or more toxins
produced by an animal.[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; 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 Wasp sting, with a droplet of venom
cause tens of thousands of human deaths per year.
However, the toxins in many venoms have potential to
treat a wide range of diseases.
Contents
Evolution
Mechanisms
Taxonomic range
Arthropods
Other invertebrates
Vertebrates
Fish
Amphibians
Reptiles
Mammals
Venom and humans
Venom Resistance
Coevolutionary Adaptations
Venom Resistance in Vertebrates
California Ground Squirrel and Northern Pacific Rattlesnake
Eels and Sea Snakes
Clownfish and Sea Anemones
Kingsnakes
See also
References
https://en.wikipedia.org/wiki/Venom 1/13
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
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]
Necrotoxins, which cause necrosis (i.e., death) in the cells

they encounter.[4] The venom of most viper species contains
phospholipase and trypsin-like serine proteases.
Neurotoxins, which primarily affect the nervous systems of
animals.[5] These include 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)
Phospholipase A2, an enzyme
and lizard venoms.[6][7][8][9] found in bee stings.
Cytotoxins, which kill individual cells, are found in the apitoxin
of honey bees and the venom of black widow spiders.[10][11]
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 — 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 Venomous animals
bugs and many ants, also produce venom.[18] At least one ant
species (Polyrhachis dives) has been shown to use venom
topically for the sterilisation of pathogens.[19]
Other invertebrates
There are venomous invertebrates in several phyla, including jellyfish such as the dangerous box
jellyfish[20] and sea anemones among the Cnidaria,[21] sea urchins among the Echinodermata,[22] and
cone snails[23] and cephalopods including octopuses among the Molluscs.[24]
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.[25]
Amphibians
Among amphibians, some salamanders can extrude sharp venom-tipped ribs.[26][27]
Reptiles
Some 450 species of snake are venomous.[25] 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
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.
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 their venom is as complex as the mixture of 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.[25][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]
Solenopsins extracted from fire ant venom has demonstrated biomedical applications, ranging from
cancer treatment to psoriasis.[46][47]
Venom Resistance
Coevolutionary Adaptations
Venom is utilized as a trophic weapon by multiple predator species. The coevolution between
predators and prey is a driving force of venom resistance, which has evolved multiple times
throughout the animal kingdom. Repeated interactions between two species can generate
coevolution.[48] The coevolution between venomous predators and venom resistant prey is best
described as a chemical arms race.[49] Predator and prey pairs are expected to associate with one
another for stable periods of time.[50] Venom is used as a chemical weapon by predator species. As the
predator capitalizes on susceptible individuals, the surviving individuals are limited to those who are
able to evade predation.[51] Resistance phenotypes typically increase over time as the predator
becomes increasingly unable to subdue prey that have developed this new resistance phenotype.[52]
The cost of developing a venom resistance is high, for predator and prey.[53] Developing an entire
physiological resistance is extremely costly, however it maximizes chances of survival for prey species
and allows predator species to expand into underutilized trophic niches. If it is possible for an animal
to evade predation through something less costly like a behavioral modification, the development of a
physiological modification becomes unnecessary.[54]
Venom Resistance in Vertebrates
California Ground Squirrel and Northern Pacific Rattlesnake

One of the most heavily researched cases of venom resistance is
the California Ground Squirrel, which is resistant to the venom of
the Northern Pacific Rattlesnake. The predator-prey pair have
coexisted for generations. The repeated interactions fostered the
development of an anti-snake venom defense in the California
ground squirrels. Researchers found evidence supporting the
theory that venom resistance is driven by coevolution among
these populations of California ground squirrels.[55] They use
toxin scavenging to negate the effects of the haemolytic toxins of Northern Pacific Rattlesnake
their rattlesnake predators, demonstrating a physiological (Crotalis viridis oraganus)
resistance to rattlesnake venom. Resistance in these ground
squirrels is population dependent. In areas where rattlesnake
populations are very dense, there is a significant increase in resistance in the squirrels compared to
populations where rattlesnakes are rare.[56] Rattlesnakes demonstrated local adaptions in the
effectiveness of their venom in order to overcome the venom resistant squirrels.[57]
Eels and Sea Snakes
There is an ongoing evolutionary contest between toxicity and resistance. The resistance of eels to sea
snake venom is a good example of coevolution between predator-prey pairs. Sea snake venom is
composed of complex mixtures of neurotoxins, myotoxins, nephrotoxins, and other nontoxic
substances.[58] The composition of the sea snake venom is species specific. The biggest piece of
evidence for this as a case of coevolution is that eels that are favored by sea snakes as prey have
unusually high tolerances to the venom of the sea snake.[59] Studies have analyzed the resistance of
four species of eels to two different sea snakes: one is a dietary generalist and the other is an eel
specialist.[60] The eels were more resistant to the venom of the eel specialist sea snake. Non-prey
fishes exhibited very low levels of resistance to the sea snake venom, further supporting
coevolution.[60]
Clownfish and Sea Anemones

Clownfish make their homes in the safety among the tentacles of sea anemones. Sea anemones are
one of the most venomous organisms on the planet, yet clownfish are able to inhabit the sea anemone
unharmed.[61] The genetic mechanisms allowing clownfish to interact with sea anemones is still
somewhat unclear.[62] Only 10 known species of anemones are hosts to clownfish and only certain
pairs of anemones and clownfish are compatible with one another.[63][64]
All sea anemones produce venoms delivered through discharging

nematocysts and mucous secretions. The toxins are composed of
peptides and proteins. They are used for prey acquisition and to
deter predators by causing pain, loss of muscular coordination
and tissue damage. Clownfish have a protective mucous that acts
as a chemical camouflage or macromolecular mimicry preventing
“not self” recognition by the sea anemone and nematocyst
discharge.[65] The sea anemone perceives the fish as its “self”
most likely by the same mechanism which prevents the discharge Ocellaris clownfish (Amphiprion
of nematocysts when its tentacles come into contact with each ocellaris) with host anemone
other. [66] Clownfish exhibit either strict host specificity or are
environmental niche generalists and can associate with a variety
of sea anemone species.[67] In some species, the mucus of clownfish was found to change during
acclimation to resemble that of the specific species of sea anemone.[67] For the Clownfish, a
relationship with the sea anemone is an obligatory one. In some cases, it is an obligatory relationship
for the anemone as well. In all cases, the interaction between the two is mutually beneficial. Clownfish
and sea anemones are one of the most compelling cases of symbiosis. The relationship provides
mutual protection from predators and the exchange of nutrients. Divergent natural selection drives
adaptive diversification through ecological speciation. The ability of clownfish to use sea anemones as
hosts has evolved 4 independent times. The obligate relationship between the clownfish and the sea
anemone has allowed the radiation of clownfishes.[68]
Kingsnakes
Inhabiting the Americas from southeastern Canada to southern Ecuador, Kingsnakes, of genus
Lampropeltis, are constrictors that prey on many venomous snakes.[69] In order to prey upon the
venomous snakes, Kingsnakes have actually evolved a resistance rather than incrementally increasing
their resistance to a point of immunity, like many species have. Kingsnakes resistance levels are
currently known to be fixed for the duration of its life and have not been found to change with age or
exposure. It is thought, that Kingsnakes have developed this evolutionary adaptation through a
process called a co-evolutionary arms race with natural selection at the forefront.[70] Kingsnake
predators that were slightly better able to tolerate the effects of the venom were more likely to survive
with resulting genocide of the Kingsnakes that were inherently not sufficiently resistant. While
simultaneously, the venomous snakes with more potent venom were more likely to survive the
predatory nature of the Kingsnakes, thus escalating the arms race.
However, the nature of the arms race has put a stipulation onto the Kingsnakes. Kingsnakes have
evolved resistance only to the venom of snakes that are in their immediate environment, like
copperheads, cottonmouths, and North American rattlesnakes, but not to the venom of, for example,
king cobras or black mambas. Even through geographical boundaries, Kingsnake venom resistance
has varied between species. They found that blood from Eastern Kingsnakes (Lampropeltis getula)
had the widest spectrum of protection against the venoms tested and was the most effective at
neutralizing many rattlesnake venoms, but the least effective against copperhead venom. Blood from
kingsnakes from Florida & the Gulf Coast was the most effective at neutralizing the venom of
copperheads & cottonmouths. Mole Kingsnake (Lampropeltis calligaster) blood is about 75% as
effective at neutralizing Mojave Rattlesnake (Crotalus scutulatus) venom as the blood of Eastern
Kingsnakes. Gray-banded Kingsnakes (L. alterna) have moderate neutralization potential against
Western Diamondback (C. atrox) venom, but none against Eastern Diamondback (C. adamanteus)
venom.[71]
See also
Poison
Envenomation
Schmidt Sting Pain Index
Big Four (Indian snakes)
List of venomous animals
Venomous mammals
Venoms in medicine
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12/14/2019 Venom - Wikipedia

Venoms kill through the action of at least four major

Necrotoxins, which cause necrosis (i.e., death) in the cells

applied all over their body as an antimicrobial protection.[15] The

Many caterpillars have defensive venom glands associated with

Bees synthesize and employ an acidic venom (apitoxin) to cause

Venom and humans

Venom Resistance in Vertebrates

California Ground Squirrel and Northern Pacific Rattlesnake

Eels and Sea Snakes

Clownfish and Sea Anemones

All sea anemones produce venoms delivered through discharging

29. "Snake Bites" (http://eclinicalworks.adam.com/content.aspx?productId=39&pid=1&gid=000031&pr

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