Astyanax mexicanus
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The Mexican tetra (''Astyanax mexicanus''), also known as the blind cave fish, blind cave characin, and blind cave tetra, is a freshwater
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...
of the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Characidae Characidae, the characids or characins is a family of freshwater subtropical and tropical fish, belonging to the order Characiformes. The name "characins" is the historical one, but scientists today tend to prefer "characids" to reflect their ...
of the order
Characiformes Characiformes is an order of ray-finned fish, comprising the characins and their allies. Grouped in 18 recognized families, more than 2000 different species are described, including the well-known piranha and tetras.; Buckup P.A.: "Relationsh ...
. The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
of its
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
, it is native to the Nearctic realm, originating in the lower Rio Grande and the Neueces and Pecos Rivers in
Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest U.S. state by ...
, as well as the central and eastern parts of Mexico. Growing to a maximum
total length Fish measurement is the measuring of individual fish and various parts of their anatomies. These data are used in many areas of ichthyology, including taxonomy and fisheries biology. Overall length * Standard length (SL) is the length of a fish ...
of , the Mexican tetra is of typical characin shape, with unremarkable, drab coloration. Its blind cave form, however, is notable for having no eyes or pigment; it has a pinkish-white color to its body (resembling an
albino Albinism is the congenital absence of melanin in an animal or plant resulting in white hair, feathers, scales and skin and pink or blue eyes. Individuals with the condition are referred to as albino. Varied use and interpretation of the term ...
). This fish, especially the blind variant, is reasonably popular among aquarists. ''A. mexicanus'' is a peaceful species that spends most of its time in midlevel water above the rocky and sandy bottoms of pools and backwaters of creeks and rivers of its native environment. Coming from a subtropical climate, it prefers water with 6.5–8 pH, a
hardness In materials science, hardness (antonym: softness) is a measure of the resistance to localized plastic deformation induced by either mechanical indentation or abrasion. In general, different materials differ in their hardness; for example hard ...
of up to 30 dGH, and a temperature range of . In the winter, some populations migrate to warmer waters. Its natural diet consists of
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...
s,
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...
s, and annelids, although in captivity it is
omnivorous An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
. The Mexican tetra has been treated as a subspecies of '' A. fasciatus'', but this is not widely accepted. Additionally, the blind cave form is sometimes recognized as a separate species, '' A. jordani'', but this directly contradicts
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
evidence.


Blind cave form

''A. mexicanus'' is famous for its blind cave form, which is known by such names as blind cave tetra, blind tetra (leading to easy confusion with the Brazilian '' Stygichthys typhlops''), blind cave characin and blind cavefish. Depending on the exact population, cave forms can have degenerated
sight Visual perception is the ability to interpret the surrounding environment through photopic vision (daytime vision), color vision, scotopic vision (night vision), and mesopic vision (twilight vision), using light in the visible spectrum reflecte ...
or have total loss of sight and even their eyes, due to down-regulation of the protein αA-crystallin and consequent lens cell death. The fish in the Pachón caves have lost their eyes completely whilst the fish from the Micos cave only have limited sight. Cave fish and surface fish are able to produce fertile offspring. These fish can still, however, find their way around by means of their lateral lines, which are highly sensitive to fluctuating
water pressure Pressure (symbol: ''p'' or ''P'') is the force applied perpendicular to the surface of an object per unit area over which that force is distributed. Gauge pressure (also spelled ''gage'' pressure)The preferred spelling varies by country and e ...
. Blindness in A. ''mexicanus'' induces a disruption of early neuromast patterning, which further causes asymmetries in cranial bone structure. One such asymmetry is a bend in the dorsal region of their skull, which is propounded to increase water flow to the opposite side of the face, functionally enhancing sensory input and spatial mapping in the dark waters of caves. Scientists suggest that gene
cystathionine beta synthase Cystathionine-β-synthase, also known as CBS, is an enzyme () that in humans is encoded by the ''CBS'' gene. It catalyzes the first step of the transsulfuration pathway, from homocysteine to cystathionine: : L-serine + L-homocysteine \rightlefth ...
-a mutation restricts blood flow to cavefish eyes during a critical stage of growth so the eyes are covered by skin. Currently, about 30 cave populations are known, dispersed over three geographically distinct areas in a karst region of San Luis Potosí and far southern
Tamaulipas Tamaulipas (), officially the Free and Sovereign State of Tamaulipas ( es, Estado Libre y Soberano de Tamaulipas), is a state in the northeast region of Mexico; one of the 31 states which, along with Mexico City, comprise the 32 Federal Entiti ...
, northeastern Mexico. Among the various cave population are at least three with only full cave forms (blind and without pigment), at least eleven with cave, "normal" and intermediate forms, and at least one with both cave and "normal" forms but no intermediates. Studies suggest at least two distinct genetic lineages occur among the blind populations, and the current distribution of populations arose by at least five independent invasions. Furthermore, cave populations have a very recent origin (< 20,000 years) in which blindness or reduced vision evolved convergently after surface ancestors populated several caves independently and at different times. This recent origin suggests that the phenotypic changes in cavefish populations, namely eye degeneration, arose as a result of the high fixation of genetic variants present in surface fish populations in a short period of time. The eyed and eyeless forms of ''A. mexicanus'', being members of the same species, are closely related and can interbreed making this species an excellent model organism for examining convergent and
parallel evolution Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and paral ...
, regressive evolution in cave animals, and the genetic basis of regressive traits. This, combined with the ease of maintaining the species in captivity, has made it the most studied cavefish and likely also the most studied cave organism overall. The blind and colorless cave form of ''A. mexicanus'' is sometimes recognized as a separate species, '' A. jordani'', but this leaves the remaining ''A. mexicanus'' as a paraphyletic species and ''A. jordani'' as
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...
. The Cueva Chica Cave in the southern part of the Sierra del Abra system is the type locality for ''A. jordani''. Other blind populations were initially also recognized as separate species, including ''antrobius'' described in 1946 from the Pachón Cave and ''hubbsi'' described in 1947 from the Los Sabinos Cave (both subsequently merged into ''jordani''/''mexicanus''). The most divergent cave population is the one in Los Sabinos. Another cave-adapted population of ''Astyanax'', varying from blind and depigmented to individuals showing intermediate features, is known from the Granadas Cave, part of the
Balsas River The Balsas River (Spanish Río Balsas, also locally known as the Mezcala River, or Atoyac River) is a major river of south-central Mexico. The basin flows through the states of Guerrero, México, Morelos, and Puebla. Downstream of Ciudad Alta ...
drainage in
Guerrero Guerrero is one of the 32 states that comprise the 32 Federal Entities of Mexico. It is divided in 81 municipalities and its capital city is Chilpancingo and its largest city is Acapulcocopied from article, GuerreroAs of 2020, Guerrero the pop ...
, southern Mexico, but it is a part of '' A. aeneus'' (itself sometimes included in ''A. mexicanus'').


Evolution research

The surface and cave forms of the Mexican tetra have proven powerful subjects for scientists studying
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
. When the surface-dwelling ancestors of current cave populations entered the subterranean environment, the change in ecological conditions rendered their
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological pr ...
—which included many biological functions dependent on the presence of light—subject to
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
and
genetic drift Genetic drift, also known as allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene variants to disappear completely and there ...
. One of the most striking changes to evolve was the loss of eyes. This is referred to as a "regressive trait" because the surface fish that originally colonized caves possessed eyes. In addition to regressive traits, cave forms evolved "constructive traits". In contrast to regressive traits, the purpose or benefit of constructive traits is generally accepted. Active research focuses on the mechanisms driving the evolution of regressive traits, such as the loss of eyes, in ''A. mexicanus''. Recent studies have produced evidence that the mechanism may be direct selection, or indirect selection through antagonistic pleiotropy, rather than genetic drift and neutral mutation, the traditionally favored hypothesis for regressive evolution. The blind form of the Mexican tetra is different from the surface-dwelling form in a number of ways, including having unpigmented skin, having a better olfactory sense by having
taste bud Taste buds contain the taste receptor cells, which are also known as gustatory cells. The taste receptors are located around the small structures known as papillae found on the upper surface of the tongue, soft palate, upper esophagus, the c ...
s all over its head, and by being able to store four times more energy as fat, allowing it to deal with irregular food supplies more effectively. Darwin said of sightless fish: Modern genetics has made clear that the lack of use does not, in itself, necessitate a feature's disappearance. In this context, the positive genetic benefits have to be considered, i.e., what advantages are obtained by cave-dwelling tetras by losing their eyes? Possible explanations include: *Not developing eyes allows the individual more energy for growth but not egg production. However the species does use other methods to locate food and detect danger, which also consume energy that would be conserved if it had eyes or transparent eyelids. *There remains less chance of accidental damage and infection, since the previously useless and exposed organ is sealed with a flap of protective skin. It is unknown why this species did not develop transparent skin or eyelids instead, as some species of reptiles did. *The lack of eyes disables the "body clock", which is controlled by periods of light and dark, conserving energy. However sunlight does have minimal impact on the "body clock" in caves. Another likely explanation for the loss of its eyes is that of selective neutrality and genetic drift; in the dark environment of the cave, the eyes are neither advantageous nor disadvantageous and thus any genetic factors that might impair the eyes (or their development) can take hold with no consequence on the individual or species. Because there is no selection pressure for sight in this environment, any number of genetic abnormalities that give rise to the damage or loss of eyes could proliferate among the population with no effect on the fitness of the population. Among some
creationists Creationism is the religious belief that nature, and aspects such as the universe, Earth, life, and humans, originated with supernatural acts of Creation myth, divine creation.#Gunn 2004, Gunn 2004, p. 9, "The ''Concise Oxford Dictionary'' say ...
, the cave tetra is seen as evidence 'against' evolution. One argument claims this is an instance of " devolution"—showing an evolutionary trend of decreasing complexity. But evolution is a non-directional process, and while increased complexity is a common effect, there is no reason why evolution cannot tend towards simplicity if that makes an organism better suited to its environment. Inhibition of the HSP90 protein has a dramatic effect in the development of the blind tetra.


In the aquarium

The blind cave tetras seen in the aquarium trade are all based on stock collected in the Cueva Chica Cave in the southern part of the Sierra del Abra system in 1936. These were sent to an aquarium company in Texas, who soon started to distribute them to aquarists. Since then, these have been selectively bred for their troglomorphic traits. Today large numbers are bred at commercial facilities, especially in Asia. The blind cave tetra is a hardy species. Their lack of sight does not hinder their ability to get food. They prefer subdued lighting with a rocky substrate, like gravel, mimicking their natural environment. They become semi-aggressive as they age, and are by nature schooling fish. Experiments have shown that keeping these fish in bright aquarium set-ups has no effect on the development of the skin flap that forms over their eyes as they grow.


See also

* List of freshwater aquarium fish species


References

{{DEFAULTSORT:Mexican Tetra Tetras Astyanax (fish) Freshwater fish of Mexico Freshwater fish of the United States Cave fish Blind animals Least concern biota of the United States Fish described in 1853 Taxa named by Filippo De Filippi