Axolotl

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Axolotl
Axolotl ganz.jpg
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Order: Urodela
Family: Ambystomatidae
Genus: Ambystoma
Species:
A. mexicanum
Binomial name
Ambystoma mexicanum
(Shaw and Nodder, 1798)
Axolotl distribution map.svg
Its distribution is marked in red.
Synonyms [2]
  • Gyrinus mexicanus Shaw and Nodder, 1798
  • Siren pisciformis Shaw, 1802
  • Siredon axolotl Wagler, 1830
  • Axolotes guttata Owen, 1844
  • Siredon Humboldtii Duméril, Bibron, and Duméril, 1854
  • Amblystoma weismanni Wiedersheim, 1879
  • Siredon edule Dugès, 1888

The axolotl ( /ˈæksəlɒtəl/ ; from Classical Nahuatl : āxōlōtl [aːˈʃoːloːtɬ] ( Loudspeaker.svg listen )), Ambystoma mexicanum, [2] is a paedomorphic salamander related to the tiger salamander. [2] [3] [4] The species was originally found in several lakes, such as Lake Xochimilco underlying Mexico City. [1] Axolotls are unusual among amphibians in that they reach adulthood without undergoing metamorphosis. Instead of taking to the land, adults remain aquatic and gilled.

Contents

Axolotls should not be confused with the larval stage of closely related tiger salamander species (A. tigrinum), which are widespread in much of North America and occasionally become paedomorphic. Neither should they be confused with mudpuppies (Necturus spp.), fully aquatic salamanders from a different family that are not closely related to the axolotl but bear a superficial resemblance. [5]

As of 2020, wild axolotls were near extinction [6] [7] due to urbanization in Mexico City and consequent water pollution, as well as the introduction of invasive species such as tilapia and perch. They are listed as critically endangered in the wild, with a decreasing population of around 50 to 1000 adult individuals, by the International Union for Conservation of Nature and Natural Resources (IUCN) and as an endangered species by the IUCN's CITES treaty. Axolotls are used extensively in scientific research due to their ability to regenerate limbs, gills and parts of their eyes and brains. [8] Axolotls were also sold as food in Mexican markets and were a staple in the Aztec diet. [9]

Description

A captive axolotl AxolotlBE.jpg
A captive axolotl
Axolotl's gills (Ambystoma mexicanum) Axolot's gills (Ambystoma mexicanum).jpg
Axolotl's gills (Ambystoma mexicanum)

A sexually mature adult axolotl, at age 18–24 months, ranges in length from 15 to 45 cm (6 to 18 in), although a size close to 23 cm (9 in) is most common and greater than 30 cm (12 in) is rare. Axolotls possess features typical of salamander larvae, including external gills and a caudal fin extending from behind the head to the vent. [10] [11] External gills are usually lost when salamander species mature into adulthood, although the Axolotl maintains this feature. [12] This is due to their neoteny evolution, where Axolotls are much more aquatic than other salamander species. [13]

Their heads are wide, and their eyes are lidless. Their limbs are underdeveloped and possess long, thin digits. Males are identified by their swollen cloacae lined with papillae, while females are noticeable for their wider bodies full of eggs. Three pairs of external gill stalks (rami) originate behind their heads and are used to move oxygenated water. The external gill rami are lined with filaments (fimbriae) to increase surface area for gas exchange. [12] Four-gill slits lined with gill rakers are hidden underneath the external gills, which prevent food from entering and allows particles to filter through.

Axolotls have barely visible vestigial teeth, which develop during metamorphosis. The primary method of feeding is by suction, during which their rakers interlock to close the gill slits. External gills are used for respiration, although buccal pumping (gulping air from the surface) may also be used to provide oxygen to their lungs. [12] Buccal pumping can occur in a two-stroke manner that pumps air from the mouth to the lungs, and with four-stroke that reverses this pathway with compression forces.

Buccal pumping Buccal pumping.jpg
Buccal pumping
Axolotls displaying variations in color Three Colors of Axolotl.jpg
Axolotls displaying variations in color

Axolotls have four pigmentation genes; when mutated they create different color variants. The normal wild-type animal is brown/tan with gold speckles and an olive undertone. The five more common mutant colors are leucistic (pale pink with black eyes), golden albino (golden with gold eyes), xanthic (grey with black eyes), albino (pale pink/white with red eyes) in which is more common in axolotls than some other creatures and melanoid (all black/dark blue with no gold speckling or olive tone). [14] In addition, there is wide individual variability in the size, frequency, and intensity of the gold speckling and at least one variant that develops a black and white piebald appearance on reaching maturity. Because pet breeders frequently cross the variant colors, double homozygous mutants are common in the pet trade, especially white/pink animals with pink eyes that are double homozygous mutants for both the albino and leucistic trait. [15] Axolotls also have some limited ability to alter their color to provide better camouflage by changing the relative size and thickness of their melanophores. [16]

Habitat and ecology

Lake Xochimilco, Mexico City (Amanecer en Xochimilco). The native habitat of Axolotls, important to the study of preservation and conservation. Sunrise in Xochimilco (Amanecer en Xochimilco) 2.jpg
Lake Xochimilco, Mexico City (Amanecer en Xochimilco). The native habitat of Axolotls, important to the study of preservation and conservation.

The axolotl is only native to the freshwater of Lake Xochimilco and Lake Chalco in the Valley of Mexico. Lake Chalco no longer exists, having been drained as a flood control measure, and Lake Xochimilco remains a remnant of its former self, existing mainly as canals. The water temperature in Xochimilco rarely rises above 20 °C (68 °F), though it may fall between 6 and 7 °C in the winter, and perhaps lower. [17]

Surveys in 1998, 2003, and 2008 found 6,000, 1,000, and 100 axolotls per square kilometer in its Lake Xochimilco habitat, respectively. [18] A four-month-long search in 2013, however, turned up no surviving individuals in the wild. Just a month later, two wild ones were spotted in a network of canals leading from Xochimilco. [19]

The wild population has been put under heavy pressure by the growth of Mexico City. The axolotl is currently on the International Union for Conservation of Nature's annual Red List of threatened species. Non-native fish, such as African tilapia and Asian carp, have also recently been introduced to the waters. These new fish have been eating the axolotls' young, as well as their primary source of food. [20]

Axolotls are members of the tiger salamander, or Ambystoma tigrinum , species complex, along with all other Mexican species of Ambystoma. Their habitat is like that of most neotenic species—a high-altitude body of water surrounded by a risky terrestrial environment. These conditions are thought to favor neoteny. However, a terrestrial population of Mexican tiger salamanders occupies and breeds in the axolotl's habitat.[ citation needed ]

The axolotl is carnivorous, consuming small prey such as molluscs, [21] worms, insects, other arthropods, [21] and small fish in the wild. Axolotls locate food by smell, and will "snap" at any potential meal, sucking the food into their stomachs with vacuum force. [22]

Use as a model organism

Axolotl in captivity Ambystoma mexicanum 1.jpg
Axolotl in captivity

Today, the axolotl is still used in research as a model organism, and large numbers are bred in captivity. They are especially easy to breed compared to other salamanders in their family, which are rarely captive-bred due to the demands of terrestrial life. One attractive feature for research is the large and easily manipulated embryo, which allows viewing of the full development of a vertebrate. Axolotls are used in heart defect studies due to the presence of a mutant gene that causes heart failure in embryos. Since the embryos survive almost to hatching with no heart function, the defect is very observable. The axolotl is also considered an ideal animal model for the study of neural tube closure due to the similarities between human and axolotl neural plate and tube formation; the axolotl's neural tube, unlike the frog's, is not hidden under a layer of superficial epithelium. [23] There are also mutations affecting other organ systems some of which are not well characterized and others that are. [24] The genetics of the color variants of the axolotl have also been widely studied. [15]

Regeneration

The feature of the axolotl that attracts most attention is its healing ability: the axolotl does not heal by scarring and is capable of the regeneration of entire lost appendages in a period of months, and, in certain cases, more vital structures, such as tail, limb, central nervous system, and tissues of the eye and heart. [25] They can even restore less vital parts of their brains. They can also readily accept transplants from other individuals, including eyes and parts of the brain—restoring these alien organs to full functionality. In some cases, axolotls have been known to repair a damaged limb, as well as regenerating an additional one, ending up with an extra appendage that makes them attractive to pet owners as a novelty. In metamorphosed individuals, however, the ability to regenerate is greatly diminished. The axolotl is therefore used as a model for the development of limbs in vertebrates. [26]

It is believed that during limb generation, axolotls have a different system to regulate their internal macrophage level and suppress inflammation, as scarring prevents proper healing and regeneration. [27] However, this belief has been questioned by other studies. [28]

Genome

The 32 billion base pair long sequence of the axolotl's genome was published in 2018 and is the largest animal genome completed so far. It revealed species-specific genetic pathways that may be responsible for limb regeneration. [29] Although the axolotl genome is about 10 times as large as the human genome, it encodes a similar number of proteins, namely 23,251 [29] (the human genome encodes about 20,000 proteins). The size difference is mostly explained by a large fraction of repetitive sequences, but such repeated elements also contribute to increased median intron sizes (22,759 bp) which are 13, 16 and 25 times that observed in human (1,750 bp), mouse (1,469 bp) and Tibetan frog (906 bp), respectively. [29]

Neoteny

Axolotls exhibit neoteny, meaning that they reach sexual maturity without undergoing metamorphosis. [30] Many species within the axolotl's genus are either entirely neotenic or have neotenic populations. In the axolotl, metamorphic failure is caused by a lack of thyroid stimulating hormone, which is used to induce the thyroid to produce thyroxine in transforming salamanders. The genes responsible for neoteny in laboratory animals may have been identified; however, they are not linked in wild populations, suggesting artificial selection is the cause of complete neoteny in laboratory and pet axolotls. [31]

Six adult axolotls (including a leucistic specimen) were shipped from Mexico City to the Jardin des Plantes in Paris in 1863. Unaware of their neoteny, Auguste Duméril was surprised when, instead of the axolotl, he found in the vivarium a new species, similar to the salamander.[ French language verification needed ] This discovery was the starting point of research about neoteny. It is not certain that Ambystoma velasci specimens were not included in the original shipment.[ citation needed ] Vilem Laufberger in Prague used thyroid hormone injections to induce an axolotl to grow into a terrestrial adult salamander. The experiment was repeated by Englishman Julian Huxley, who was unaware the experiment had already been done, using ground thyroids. [32] Since then, experiments have been done often with injections of iodine or various thyroid hormones used to induce metamorphosis. [13]

Neoteny has been observed in all salamander families in which it seems to be a survival mechanism, in aquatic environments only of mountain and hill, with little food and, in particular, with little iodine. In this way, salamanders can reproduce and survive in the form of a smaller larval stage, which is aquatic and requires a lower quality and quantity of food compared to the big adult, which is terrestrial. If the salamander larvae ingest a sufficient amount of iodine, directly or indirectly through cannibalism, they quickly begin metamorphosis and transform into bigger terrestrial adults, with higher dietary requirements. [33] In fact, in some high mountain lakes there live dwarf forms of salmonids that are caused by deficiencies in food and, in particular, iodine, which causes cretinism and dwarfism due to hypothyroidism, as it does in humans.

Unlike some other neotenic salamanders (sirens and Necturus ), axolotls can be induced to metamorphose by an injection of iodine (used in the production of thyroid hormones) or by shots of thyroxine hormone. The adult form resembles a terrestrial plateau tiger salamander, but has several differences, such as longer toes, which support its status as a separate species.[ citation needed ]

Captive care

These axolotls at Vancouver Aquarium are leucistic, with less pigmentation than normal. Ambystoma mexicanum at Vancouver Aquarium.jpg
These axolotls at Vancouver Aquarium are leucistic, with less pigmentation than normal.

The axolotl is a popular exotic pet like its relative, the tiger salamander (Ambystoma tigrinum). As for all poikilothermic organisms, lower temperatures result in slower metabolism and a very unhealthily reduced appetite. Temperatures at approximately 16 °C (61 °F) to 18 °C (64 °F) are suggested for captive axolotls to ensure sufficient food intake; stress resulting from more than a day's exposure to lower temperatures may quickly lead to disease and death, and temperatures higher than 24 °C (75 °F) may lead to metabolic rate increase, also causing stress and eventually death. [34] [35] Chlorine, commonly added to tapwater, is harmful to axolotls. A single axolotl typically requires a 150-litre (40-US-gallon) tank. Axolotls spend the majority of the time at the bottom of the tank. [36]

This animal was x-rayed several times as part of a research project over a period of two years. It was a normal healthy adult (26.3 cm; 159.5 gm) at the beginning of the project and lived several more years after the project ended. Axolotl with Grit XRay.png
This animal was x-rayed several times as part of a research project over a period of two years. It was a normal healthy adult (26.3 cm; 159.5 gm) at the beginning of the project and lived several more years after the project ended.

Salts, such as Holtfreter's solution, are usually added to the water to prevent infection. [38]

In captivity, axolotls eat a variety of readily available foods, including trout and salmon pellets, frozen or live bloodworms, earthworms, and waxworms. Axolotls can also eat feeder fish, but care should be taken as fish may contain parasites. [39]

Substrates are another important consideration for captive axolotls, as axolotls (like other amphibians and reptiles) tend to ingest bedding material together with food [40] and are commonly prone to gastrointestinal obstruction and foreign body ingestion. [41] Some common substrates used for animal enclosures can be harmful for amphibians and reptiles. If gravel (common in aquarium use) is used, it is recommended that it consist of smooth particles of a size small enough to pass through the digestive tract. [40] One guide to axolotl care for laboratories notes that bowel obstructions are a common cause of death, and recommends that no items with a diameter below 3 cm should be available to the animal. [42]

There is some evidence that axolotls might seek out appropriately-sized gravel for use as gastroliths [43] based on experiments conducted at the University of Manitoba axolotl colony. [44] [45]

Axolotl is one of several words, mostly of foreign origin, adopted by Mad Magazine as nonsense words for use as running gags; potrzebie and veeblefetzer are two others.

In Frank Herbert's fictional Dune universe, the Tleilaxu use a technology called "axlotl tanks" to regrow limbs.

Argentine writer Julio Cortázar included a short story entitled "Axolotl" in his 1956 collection Final del juego . The story concerns a man who becomes obsessed with the salamanders after viewing them in an aquarium in Paris. [46]

In the Netflix series BoJack Horseman , there is an anthropomorphic axolotl character named Yolanda Buenaventura voiced by Natalie Morales. [47]

In 2020, it was announced that the axolotl will be featured on the new design for Mexico's 50-peso banknote, along with images of maize and chinampas. The banknotes are expected to go into circulation by 2022. [48]

Mojang Studios added Axolotls to their sandbox game, Minecraft, in update 1.17 to increase awareness of their endangerment.

See also

Related Research Articles

Amphibian Class of ectothermic tetrapods

Amphibians are ectothermic, tetrapod vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arboreal or freshwater aquatic ecosystems. Thus amphibians typically start out as larvae living in water, but some species have developed behavioural adaptations to bypass this.

Neoteny, also called juvenilization, is the delaying or slowing of the physiological development of an organism, typically an animal. Neoteny is found in modern humans. In progenesis, sexual development is accelerated.

Salamander Order of amphibians

Salamanders are a group of amphibians typically characterized by a lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All 10 present-day salamander families are grouped together under the order Urodela. Salamander diversity is highest in the Northern Hemisphere and most species are found in the Holarctic realm, with some species present in the Neotropical realm.

Vertebrate Subphylum of chordates with backbones

Vertebrates comprise all species of animals within the subphylum Vertebrata. Vertebrates represent the overwhelming majority of the phylum Chordata, with currently about 69,963 species described. Vertebrates comprise such groups as the following:

Mole salamander Genus of amphibians

The mole salamanders are a group of advanced salamanders endemic to North America. The group has become famous due to the presence of the axolotl, widely used in research due to its paedomorphosis, and the tiger salamander which is the official amphibian of many states, and often sold as a pet.

Tiger salamander Species of amphibian

The tiger salamander is a species of mole salamander and one of the largest terrestrial salamanders in North America.

Spotted salamander Species of amphibian

The spotted salamander or yellow-spotted salamander is a mole salamander common in eastern United States and Canada. The spotted salamander is the state amphibian of Ohio and South Carolina. This salamander ranges from Nova Scotia, to Lake Superior, to southern Georgia and Texas. Its embryos have been found to have symbiotic algae living inside them, the only known example of vertebrate cells hosting an endosymbiont microbe.

Regeneration (biology) Biological process of renewal, restoration, and tissue growth

In biology, regeneration is the process of renewal, restoration, and tissue growth that makes genomes, cells, organisms, and ecosystems resilient to natural fluctuations or events that cause disturbance or damage. Every species is capable of regeneration, from bacteria to humans. Regeneration can either be complete where the new tissue is the same as the lost tissue, or incomplete where after the necrotic tissue comes fibrosis.

Heterochrony Evolutionary change in the rates or durations of developmental events, leading to structural changes

In evolutionary developmental biology, heterochrony is any genetically controlled difference in the timing, rate, or duration of a developmental process in an organism compared to its ancestors or other organisms. This leads to changes in the size, shape, characteristics and even presence of certain organs and features. It is contrasted with heterotopy, a change in spatial positioning of some process in the embryo, which can also create morphological innovation. Heterochrony can be divided into intraspecific heterochrony, variation within a species, and interspecific heterochrony, phylogenetic variation, i.e. variation of a descendant species with respect to an ancestral species.

Common mudpuppy Species of salamander

The common mudpuppy is a species of salamander in the genus Necturus. They live an entirely aquatic lifestyle in parts of North America in lakes, rivers, and ponds. They go through paedomorphosis and retain their external gills. Because skin and lung respiration alone is not sufficient for gas exchange, mudpuppies must rely on external gills as their primary means of gas exchange. They are usually a rusty brown color and can grow to an average length of 13 in (330 mm). Mudpuppies are nocturnal creatures, and come out during the day only if the water in which they live is murky. Their diet consists of almost anything they can get in their mouths, including insects, mollusks, and earthworms. Once a female mudpuppy reaches sexual maturity at six years of age, she can lay an average of 60 eggs. In the wild, the average lifespan of a mudpuppy is 11 years.

Blastema Mass of cells capable of enacting growth and regeneration

A blastema is a mass of cells capable of growth and regeneration into organs or body parts. The changing definition of the word blastema since its introduction into the biomedical vocabulary in 1799 has been reviewed by Holland (2021). Historically, blastemas were thought to be composed of undifferentiated pluripotent cells, but recent research indicates that in some organisms blastemas may retain memory of tissue origin. Blastemas are typically found in the early stages of an organism's development such as in embryos, and in the regeneration of tissues, organs and bone.

Long-toed salamander Species of amphibian

The long-toed salamander is a mole salamander in the family Ambystomatidae. This species, typically 4.1–8.9 cm (1.6–3.5 in) long when mature, is characterized by its mottled black, brown, and yellow pigmentation, and its long outer fourth toe on the hind limbs. Analysis of fossil records, genetics, and biogeography suggest A. macrodactylum and A. laterale are descended from a common ancestor that gained access to the western Cordillera with the loss of the mid-continental seaway toward the Paleocene.

Taylors salamander Species of amphibian

Taylor's salamander, Ambystoma taylori, is a species of salamander found only in Laguna Alchichica, a high-altitude crater lake to the southwest of Perote, Puebla, Mexico. It was first described in 1982 but had been known to science prior to that. It is a neotenic salamander, breeding while still in the larval state and not undergoing metamorphosis. The lake in which it lives is becoming increasingly saline and less suitable for the salamander, which is declining in numbers. The International Union for Conservation of Nature (IUCN) has rated it as being "critically endangered".

Andersons salamander Species of amphibian

Anderson's salamander is a neotenic salamander from Zacapu Lagoon in the Mexican state of Michoacán.

The Lake Patzcuaro salamander, locally known as achoque, Ambystoma dumerilii, is a neotenic salamander species.

The Puerto Hondo stream salamander or Michoacan stream salamander, Ambystoma ordinarium, is a mole salamander from the Cordillera Volcánica within the Mexican state of Michoacán.

Yellow-peppered salamander Species of amphibian

The yellow-peppered salamander also known as the salamandra de Champala and the yellow-headed salamander, is a species of mole salamander native to areas at an elevation of 4900 ft around Santa Cruz, Rancho Malveste and Tapalpa in Jalisco, Mexico.

<i>Gerrothorax</i> Extinct genus of amphibians

Gerrothorax is an extinct genus of temnospondyl amphibian from the Triassic period of Greenland, Germany, Sweden, and possibly Thailand. It is known from a single species, G. pulcherrimus, although several other species such as G. pustuloglomeratus have been named in the past.

Branchiosauridae Extinct family of amphibians

Branchiosauridae is an extinct family of small amphibamiform temnospondyls with external gills and an overall juvenile appearance. The family has been characterized by hundreds of well-preserved specimens from the Permo-Carboniferous of Middle Europe. Specimens represent well defined ontogenetic stages and thus the taxon has been described to display paedomorphy (perennibranchiate). However, more recent work has revealed branchiosaurid taxa that display metamorphosing trajectories. The name Branchiosauridae refers to the retention of gills.

Parthenogenesis is a form of reproduction where eggs develop without fertilization, resulting in unisexual species. This phenomenon is closely related with reproductive modes such as hybridogenesis, where fertilization occurs, but the paternal DNA is not passed on. Among amphibians, it is seen in numerous frog and salamander species, but has not been recorded in caecilians.

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