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Asexual reproduction Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the fu ...
in starfish takes place by fission or through
autotomy Autotomy (from the Greek language, Greek ''auto-'', "self-" and ''tome'', "severing", wikt:αὐτοτομία, αὐτοτομία) or self-amputation, is the behaviour whereby an animal sheds or discards one or more of its own appendages, usual ...
of arms. In fission, the central disc breaks into two pieces and each portion then regenerates the missing parts. In autotomy, an arm is shed with part of the central disc attached, which continues to live independently as a "comet", eventually growing a new set of arms. Although almost all sea stars can regenerate their limbs, only a select few sea star species are able to reproduce in these ways.


Fission

Fissiparity in the starfish family
Asteriidae The Asteriidae are a diverse family of Asteroidea (sea stars) in the order Forcipulatida. It is one of three families in the order Forcipulatida. Genera The World Register of Marine Species lists these genera within the family Asteriidae (in a ...
is confined to the genera ''
Coscinasterias ''Coscinasterias'' is a genus of sea stars of the family Asteriidae. Species There are four recognized species: * '' Coscinasterias acutispina'' (Stimpson, 1862) * ''Coscinasterias calamaria'' (Gray, 1840) – Eleven-arm sea star * '' Coscinast ...
'', '' Stephanasterias'' and ''
Sclerasterias ''Sclerasterias'' is a genus of starfish in the family (biology), family Asteriidae. Adult individuals have five arms but small, immature individuals have six. This led to the giving of a separate generic name to the juveniles, ''Hydrasterias' ...
''. Another family in which asexual reproduction by fission has independently arisen is the
Asterinidae The Asterinidae are a large family of sea stars in the order Valvatida. Description and characteristics These are generally small sea stars, flattened dorsally and bearing very short arms, often giving a pentagonal shape in the body ;example: ...
. The life span is at least four years. A dense population of '' Stephanasterias albula'' was studied at North
Lubec, Maine Lubec ( ) is a town in Washington County, Maine, United States. It is the easternmost municipality in the contiguous U.S. (see extreme points of the United States) and is the country's closest continental location to Africa. The town, with a p ...
. All the individuals were fairly small, with arm lengths not exceeding , but no juveniles were found, suggesting that there had been no recent larval recruitment and that this species may be obligately fissiparous. Fission seemed to take place only in the spring and summer and for any individual, occurred once a year or once every two years. Another species, ''
Coscinasterias tenuispina ''Coscinasterias tenuispina'' is a starfish in the family Asteriidae. It is sometimes called the blue spiny starfish or the white starfish. It occurs in shallow waters in the Atlantic Ocean and the Mediterranean Sea. Description ''Coscinasteria ...
'', has a variable number of arms but is often found with 7 arms divided into dis-similar sized groups of 3 and 4. It is unclear why fission starts in any particular part of the disc rather than any other, but the origin seemed to bear some relation to the position of the
madreporite The madreporite is a light colored calcareous opening used to filter water into the water vascular system of echinoderms. It acts like a pressure-equalizing valve. It is visible as a small red or yellow button-like structure, looking like a smal ...
s and the longest arm. This species typically reproduces sexually in the winter and by fission at other times of year. The undivided individual has 1 to 5 madreporites and at least one is found in each offspring. New arms usually appear in groups of 4 and are normally accompanied by the appearance of additional madreporites. The presence of multiple madreporites seems to be a prerequisite of fission. In Brazil, only male individuals have been found and fission takes place all the year round, though primarily in the winter. Fission seems to be correlated with certain stress factors such as particularly low tides, when many starfish may be exposed to the air. ''
Nepanthia belcheri ''Nepanthia belcheri'' is a species of starfish in the family Asterinidae. It is found in shallow water in Southeast Asia and northeastern Australia. It is an unusual species in that it can reproduce sexually or can split in two by fission to fo ...
'' has a variable number of arms and divides by fission in a similar manner. It is a
hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...
, some individuals having gonads that function as testes and others gonads that function as ovaries. After fission, the gonads regress and individuals that previously had mature female gonads become masculinized, developing male-type gonads. Many larger individuals provide evidence from the varying lengths of their arms that they have divided by fission on several occasions. In ''
Sclerasterias ''Sclerasterias'' is a genus of starfish in the family (biology), family Asteriidae. Adult individuals have five arms but small, immature individuals have six. This led to the giving of a separate generic name to the juveniles, ''Hydrasterias' ...
'', fissiparity seems to be confined to very young individuals. In these, there is a transitory hexamerous symmetry in what is a normally a pentamerously symmetrical genus. The immature individuals with 6 arms appear so different in appearance from mature individuals with 5 arms that they were at one time considered to be two genera, ''Hydrasterias'' and ''Sclerasterias''. Juveniles with arms measuring between 8 mm and 15 mm (occasionally 20 mm) are usually involved in fission and undergo multiple divisions. A sample of 36 young ''
Sclerasterias euplecta ''Sclerasterias'' is a genus of starfish in the family Asteriidae. Adult individuals have five arms but small, immature individuals have six. This led to the giving of a separate generic name to the juveniles, ''Hydrasterias'', before it was r ...
'' of this size was examined. 9 had only 5 arms and did not show evidence of fissiparity while the remainder had 6 arms, usually 3 longer than the other 3, following prior fission. In another sample of juvenile '' Scierasterias heteropau'', the arms were similarly arranged in groups of three and there were 4 madreporites, 2 on the original and 2 on the regenerated section. Active fissiparity seems to be correlated with 6 arms and 4 madreporites. At some stage in their development as yet unexplained, only 5 arms and one madreporite appear, and the ability to divide in this way is lost.


Autotomy as a means of asexual reproduction


History

Writing in 1872, Lutken suggested that in certain members of the
Ophiuroidea Brittle stars, serpent stars, or ophiuroids (; ; referring to the serpent-like arms of the brittle star) are echinoderms in the class Ophiuroidea, closely related to starfish. They crawl across the sea floor using their flexible arms for locomot ...
, "a radiary division occurs in which cast off arms formed new rays and a disk". Six years later
Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new sp ...
observed that members of the genus ''Ophidiaster'' (''
Linckia ''Linckia'' is a genus of sea stars found mainly in the Indo-Pacific region. They are known to be creatures with remarkable regenerative abilities, and capable of defensive autotomy against predators. They reproduce asexually. The genus is name ...
'') were prone to cast off arms and that new discs, arms, madreporites and mouths formed on the severed surface of these. In 1904, Kellogg observed numerous severed arms on reefs at
Apia Apia () is the Capital (political), capital and largest city of Samoa, as well as the nation's only city. It is located on the central north coast of Upolu, Samoa's second-largest island. Apia falls within the political district (''itūmālō ...
,
Samoa Samoa, officially the Independent State of Samoa; sm, Sāmoa, and until 1997 known as Western Samoa, is a Polynesian island country consisting of two main islands (Savai'i and Upolu); two smaller, inhabited islands (Manono Island, Manono an ...
, noting that many were sprouting new arms and suggested that '' Linckia diplax'' and '' Linckia pacifica'' had the ability to generate new individuals in this way. He thought the arms might be shed by
autotomy Autotomy (from the Greek language, Greek ''auto-'', "self-" and ''tome'', "severing", wikt:αὐτοτομία, αὐτοτομία) or self-amputation, is the behaviour whereby an animal sheds or discards one or more of its own appendages, usual ...
. In the same year, Monks showed experimentally that the "comets" developing from the severed arms of ''
Linckia columbiae ''Linckia columbiae'' is a species of starfish in the family Ophidiasteridae. It is found in the East Pacific where it ranges from California (USA) to northwest Peru, including offshore islands such as the Galápagos.James, M.J., eds. (1991). '' ...
'' could indeed grow into new individuals.


Autotomy of arms

''
Linckia multifora ''Linckia multifora'' is a variously colored starfish in the Family (biology), family Ophidiasteridae that is found in the Indian Ocean and Red Sea. Its common names include the Dalmatian Linckia, mottled Linckia, spotted Linckia, multicolor sea ...
'' and ''
Linckia guildingi ''Linckia guildingi'', also called the common comet star, Guilding's sea star or the green Linckia, is a species of sea star reported from the shallow waters of the tropical Pacific Ocean, Indian Ocean, Atlantic Ocean and the Caribbean Sea. Taxo ...
'' are two species of starfish found on Hawaii which were found to exhibit
autotomy Autotomy (from the Greek language, Greek ''auto-'', "self-" and ''tome'', "severing", wikt:αὐτοτομία, αὐτοτομία) or self-amputation, is the behaviour whereby an animal sheds or discards one or more of its own appendages, usual ...
, shedding one or more arms frequently. The arms are known as "comets" and can move about independently and each one can grow into a new individual. Though severed from the nervous system and the
water vascular system The water vascular system is a hydraulic system used by echinoderms, such as sea stars and sea urchins, for locomotion, food and waste transportation, and respiration. The system is composed of canals connecting numerous tube feet. Echinoderms mov ...
they still exhibit normal behaviour patterns. In a study undertaken in Hawaii, it was found that the detachment of an arm was not a sudden event. Most fractures took place about from the disk and started with a small crack appearing on the lower surface of the arm. This spread laterally and upwards towards the dorsal surface. Then the
tube feet Tube feet (technically podia) are small active tubular projections on the oral face of an echinoderm, whether the arms of a starfish, or the undersides of sea urchins, sand dollars and sea cucumbers; they are more discreet though present on britt ...
on the arm and those on the body pulled the two parts of the animal in opposite directions until they parted. The process could take about an hour to complete. The damaged tissue healed in about 10 days and the animal grew a new arm over the course of several months. Breaks took place in various positions on the arm, though Crozier noted a particular breaking zone in ''
Coscinasterias tenuispina ''Coscinasterias tenuispina'' is a starfish in the family Asteriidae. It is sometimes called the blue spiny starfish or the white starfish. It occurs in shallow waters in the Atlantic Ocean and the Mediterranean Sea. Description ''Coscinasteria ...
''. The immediate cause of the autotomy is not always apparent. Of 50 specimens of ''Linckia multifora'' brought to the laboratory, 18 had shed one or more arms within 24 hours. The mortality rate of newly severed arms was high, many succumbing to bacterial infection while the wounds were fresh. Once the wound had healed, in about 10 days, survival was more likely. When arms were severed into several lengths in the laboratory, it was found that those over in length were capable of regenerating. These included the tips of the arms and the central sections with wounds at each end. It takes about 10 months to regenerate a new disk with arms in length. The first development in the regeneration cycle is the formation of a crescent-shaped ridge at the damaged end. Grooves begin to form and a mouth develops at the point from which they radiate. The arms start to form and tube feet begin to appear. As the arms grow the disc begins to develop and eventually a madreporite appears. This process lasts for some time, and about 10 months after separation, the comet has a half disc and 4 arms about long.


References

{{reflist, 30em Echinoderm biology Asexual reproduction in animals