Thelyotokous
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Thelytoky (from the Greek ''thēlys'' "female" and ''tokos'' "birth") is a type of
parthenogenesis Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...
in which females are produced from unfertilized eggs, as for example in aphids. Thelytokous parthenogenesis is rare among animals and reported in about 1,500
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, about 1 in 1000 of described animal species, according to a 1984 study. It is more common in
invertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
, like
arthropods Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
, but it can occur in
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
, including
salamander Salamanders are a group of amphibians typically characterized by their 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 ten ...
s,
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 li ...
, and
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s such as some whiptail lizards. Thelytoky can occur by different mechanisms, each of which has a different impact on the level of homozygosity. It is found in several groups of
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
, including Apidae,
Aphelinidae The Aphelinidae are a moderate-sized family of tiny parasitic wasps, with about 1100 described species in some 28 genera. These minute insects are challenging to study, as they deteriorate rapidly after death unless extreme care is taken (e.g., p ...
,
Cynipidae Gall wasps, also incorrectly called gallflies, are hymenopterans of the family Cynipidae in the wasp superfamily Cynipoidea. Their common name comes from the galls they induce on plants for larval development. About 1,300 species of this genera ...
, Formicidae, Ichneumonidae, and Tenthredinidae. It can be induced in Hymenoptera by the
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
'' Wolbachia'' and ''Cardinium''.


Arrhenotoky and thelytoky in Hymenoptera

Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
( ants,
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...
s,
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s, and sawflies) have a
haplodiploid sex-determination system Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky. Haplodiploidy determines the sex ...
. They produce haploid males from unfertilized eggs (
arrhenotoky Arrhenotoky (from Greek -τόκος ''-tókos'' "birth of -" + ἄρρην ''árrhēn'' "male person"), also known as arrhenotokous parthenogenesis, is a form of parthenogenesis in which unfertilized eggs develop into males. In most cases, parthen ...
), a form of
parthenogenesis Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...
. However, in a few social hymenopterans,
queen Queen or QUEEN may refer to: Monarchy * Queen regnant, a female monarch of a Kingdom ** List of queens regnant * Queen consort, the wife of a reigning king * Queen dowager, the widow of a king * Queen mother, a queen dowager who is the mother ...
s or workers are capable of producing diploid female offspring by thelytoky. The daughters produced may or may not be complete clones of their mother depending on the type of parthenogenesis that takes place. The offspring can develop into either queens or workers. Examples of such species include the Cape bee, ''
Apis mellifera capensis The Cape honey bee or Cape bee (''Apis mellifera capensis'') is a southern South African subspecies of the western honey bee. They play a major role in South African agriculture and the economy of the Western Cape by pollinating crops and produc ...
'', ''
Mycocepurus smithii ''Mycocepurus smithii'' is a species of fungus-growing ant from Latin America. This species is widely distributed geographically and can be found from Mexico in the north to Argentina in the south, as well as on some Caribbean Islands. It lives ...
'' and clonal raider ant, ''
Ooceraea biroi ''Ooceraea biroi'', the clonal raider ant, is a queenless clonal ant in the genus ''Ooceraea'' (recently transferred from the genus ''Cerapachys''). Native to the Asian mainland, this species has become invasive on tropical and subtropical island ...
''.


Automixis

Automixis is a form of thelytoky. In automictic
parthenogenesis Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...
,
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
takes place and diploidy is restored by fusion of first division non-sister nuclei (central fusion) or the second division sister nuclei (terminal fusion). (see diagram).


With central fusion

Automixis with central fusion tends to maintain heterozygosity in the passage of the genome from mother to daughter. This form of automixis has been observed in several ant species including the desert ant ''
Cataglyphis cursor ''Cataglyphis cursor'' is a species of ant in the genus ''Cataglyphis''. Described in 1846, it is known only from the Mediterranean parts of France. Reproduction Parthenogenesis is a natural form of reproduction in which growth and development ...
'', the clonal raider ant ''
Cerapachys biroi ''Ooceraea biroi'', the clonal raider ant, is a queenless clonal ant in the genus ''Ooceraea'' (recently transferred from the genus ''Cerapachys''). Native to the Asian mainland, this species has become invasive on tropical and subtropical islan ...
'', the predaceous ant ''
Platythyrea punctata ''Platythyrea punctata'' is a species of ant in the family Formicidae. References Further reading * Ponerinae Articles created by Qbugbot Insects described in 1858 {{ponerinae-stub ...
'', and the electric ant (little fire ant) '' Wasmannia auropunctata''. Automixis with central fusion also occurs in the Cape honey bee ''
Apis mellifera capensis The Cape honey bee or Cape bee (''Apis mellifera capensis'') is a southern South African subspecies of the western honey bee. They play a major role in South African agriculture and the economy of the Western Cape by pollinating crops and produc ...
'', the brine shrimp ''Artemia parthenogenetica'', and the termite ''Embiratermes neotenicus''.
Oocyte An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female ...
s that undergo automixis with central fusion often display a reduced rate of crossover recombination. A low rate of recombination in automictic oocytes favors maintenance of
heterozygosity Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
, and only a slow transition from heterozygosity to homozygosity over successive generations. This allows avoidance of immediate
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. In ...
. Species that display central fusion with reduced recombination include the ants ''P. punctata'' and ''W. auropunctata'', the brine shrimp ''A. parthenogenetica'', and the honey bee ''A. m. capensis''. In ''A. m. capensis'', the recombination rate during the meiosis associated with thelytokous parthenogenesis is reduced by more than 10-fold. In ''W. auropunctata'' the reduction is 45-fold. Single queen colonies of the narrow headed ant '' Formica exsecta'' provide an illustrative example of the possible deleterious effects of increased homozygosity. In this ant the level of queen homozygosity is negatively associated with colony age. Reduced colony survival appears to be due to decreased queen lifespan resulting from queen homozygosity and expression of deleterious recessive mutations (
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. In ...
).


With terminal fusion

Automixis with terminal fusion tends to promote homozygosity in the passage of the genome from mother to daughter. This form of automixis has been observed in the water flea ''
Daphnia magna ''Daphnia magna'' is a small planktonic crustacean (adult length 1.5–5.0 mm) that belongs to the subclass Phyllopoda. Description ''D. magna'' is a typical water flea of the genus ''Daphnia''. The females reach up to 5 mm in size, ...
'' and the Colombian rainbow boa constrictor ''
Epicrates maurus ''Epicrates maurus'' is a species of non-venomous constrictor in the family Boidae, commonly found in the Amazon region of South America. The common name for this species is the brown rainbow boa. This species is semi-arboreal, spending time both ...
''. Parthenogenesis in ''E. maurus'' is only the third genetically confirmed case of consecutive virgin births of viable offspring from a single female within any vertebrate lineage. However, survival of offspring over two successive litters was poor, suggesting that automixis with terminal fusion leads to homozygosity and expression of deleterious
recessive allele In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and t ...
s (
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. In ...
).


See also

*
Autotoky Autotoky is uniparental reproduction by self-fertilization or by parthenogenesis Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, labe ...
*
Epitoky Epitoky is a process that occurs in many species of polychaete marine worms wherein a sexually immature worm (the atoke) is modified or transformed into a sexually mature worm (the epitoke). Epitokes are pelagic morphs capable of sexual reproduc ...
*
Arrhenotoky Arrhenotoky (from Greek -τόκος ''-tókos'' "birth of -" + ἄρρην ''árrhēn'' "male person"), also known as arrhenotokous parthenogenesis, is a form of parthenogenesis in which unfertilized eggs develop into males. In most cases, parthen ...


References

{{Eusociality Beekeeping Insect reproduction Asexual reproduction