Parthenogenesis (/ˌpɑːrθɪnoʊˈdʒɛnɪsɪs, -θɪnə-/;[1][2] from the Greek παρθένος, parthenos, 'virgin' + γένεσις, genesis, 'creation'[3]) is a natural form of asexual reproduction in which growth and development of embryos occur without fertilization. In animals, parthenogenesis means development of an embryo from an unfertilized egg cell. In plants parthenogenesis is a component process of apomixis.
Parthenogenesis occurs naturally in some plants, some invertebrate animal species (including nematodes, water fleas, some scorpions, aphids, some mites, some bees, some Phasmida and parasitic wasps) and a few vertebrates (such as some fish,[4] amphibians, reptiles[5][6] and very rarely birds[7]).  This type of reproduction has been induced artificially in a few species including fish and amphibians.[8]
Normal egg cells form after meiosis and are haploid, with half as many chromosomes as their mother's body cells. Haploid individuals, however, are usually non-viable, and parthenogenetic offspring usually have the diploid chromosome number. Depending on the mechanism involved in restoring the diploid number of chromosomes, parthenogenetic offspring may have anywhere between all and half of the mother's alleles. The offspring having all of the mother's genetic material are called full clones and those having only half are called half clones. Full clones are usually formed without meiosis. If meiosis occurs, the offspring will get only a fraction of the mother's alleles since crossing over of DNA takes place during meiosis, creating variation.
Parthenogenetic offspring in species that use either the XY or the X0 sex-determination system have two X chromosomes and are female. In species that use the ZW sex-determination system, they have either two Z chromosomes (male) or two W chromosomes (mostly non-viable but rarely a female), or they could have one Z and one W chromosome (female).
Life history types

Some species reproduce exclusively by parthenogenesis (such as the Bdelloid rotifers), while others can switch between sexual reproduction and parthenogenesis. This is called facultative parthenogenesis (other terms are cyclical parthenogenesis, heterogamy[9][10] or heterogony[11][12]). The switch between sexuality and parthenogenesis in such species may be triggered by the season (aphid, some gall wasps), or by a lack of males or by conditions that favour rapid population growth (rotifers and cladocerans like daphnia). In these species asexual reproduction occurs either in summer (aphids) or as long as conditions are favourable. This is because in asexual reproduction a successful genotype can spread quickly without being modified by sex or wasting resources on male offspring who won't give birth. In times of stress, offspring produced by sexual reproduction may be fitter as they have new, possibly beneficial gene combinations. In addition, sexual reproduction provides the benefit of meiotic recombination between non-sister chromosomes, a process associated with repair of DNA double-strand breaks and other DNA damages that may be induced by stressful conditions.[13][14] (See also: Origin and function of meiosis.)
Many taxa with heterogony have within them species that have lost the sexual phase and are now completely asexual. Many other cases of obligate parthenogenesis (or gynogenesis) are found among polyploids and hybrids where the chromosomes cannot pair for meiosis.
The production of female offspring by parthenogenesis is referred to as thelytoky (e.g., aphids) while the production of males by parthenogenesis is referred to as arrhenotoky (e.g., bees). When unfertilized eggs develop into both males and females, the phenomenon is called deuterotoky.[15]
Types and mechanisms
Parthenogenesis can occur without meiosis through mitotic oogenesis. This is called apomictic parthenogenesis. Mature egg cells are produced by mitotic divisions, and these cells directly develop into embryos. In flowering plants, cells of the gametophyte can undergo this process. The offspring produced by apomictic parthenogenesis are full clones of their mother. Examples include aphids.
Parthenogenesis involving meiosis is more complicated. In some cases, the offspring are haploid (e.g., male ants). In other cases, collectively called automictic parthenogenesis, the ploidy is restored to diploidy by various means. This is because haploid individuals are not viable in most species.  In automictic parthenogenesis, the offspring differ from one another and from their mother. They are called half clones of their mother.
Automictic

Automixis[16] is a term that covers several reproductive mechanisms, some of which are parthenogenetic.[17]
Diploidy might be restored by the doubling of the chromosomes without cell division before meiosis begins or after meiosis is completed. This is referred to as an endomitotic cycle. This may also happen by the fusion of the first two blastomeres. Other species restore their ploidy by the fusion of the meiotic products. The chromosomes may not separate at one of the two anaphases (called restitutional meiosis) or the nuclei produced may fuse or one of the polar bodies may fuse with the egg cell at some stage during its maturation.
Some authors consider all forms of automixis sexual as they involve recombination. Many others classify the endomitotic variants as asexual and consider the resulting embryos parthenogenetic. Among these authors, the threshold for classifying automixis as a sexual process depends on when the products of anaphase I or of anaphase II are joined together. The criterion for "sexuality" varies from all cases of restitutional meiosis,[18] to those where the nuclei fuse or to only those where gametes are mature at the time of fusion.[17] Those cases of automixis that are classified as sexual reproduction are compared to self-fertilization in their mechanism and consequences.
The genetic composition of the offspring depends on what type of apomixis takes place. When endomitosis occurs before meiosis[19][20] or when central fusion occurs (restitutional meiosis of anaphase I or the fusion of its products), the offspring get all[19][21] to more than half of the mother's genetic material and heterozygosity is mostly preserved[22] (if the mother has two alleles for a locus, it is likely that the offspring will get both). This is because in anaphase I the homologous chromosomes are separated. Heterozygosity is not completely preserved when crossing over occurs in central fusion.[23] In the case of pre-meiotic doubling, recombination -if it happens- occurs between identical sister chromatids.[19]
If terminal fusion (restitutional meiosis of anaphase II or the fusion of its products) occurs, a little over half the mother's genetic material is present in the offspring and the offspring are mostly homozygous.[24] This is because at anaphase II the sister chromatids are separated and whatever heterozygosity is present is due to crossing over. In the case of endomitosis after meiosis, the offspring is completely homozygous and has only half the mother's genetic material.
This can result in parthenogenetic offspring being unique from each other and from their mother.
Sex of the offspring
In apomictic parthenogenesis, the offspring are clones of the mother and hence (except for aphids) are usually female. In the case of aphids, parthenogenetically produced males and females are clones of their mother except that the males lack one of the X chromosomes (XO).[25]
When meiosis is involved, the sex of the offspring will depend on the type of sex determination system and the type of apomixis. In species that use the XY sex-determination system, parthenogenetic offspring will have two X chromosomes and are female. In species that use the ZW sex-determination system the offspring genotype may be one of ZW (female),[21][22] ZZ (male), or WW (non-viable in most species[24] but a fertile, viable female in a few (e.g., boas)).[24] ZW offspring are produced by endoreplication before meiosis or by central fusion.[21][22] ZZ and WW offspring occur either by terminal fusion[24] or by endomitosis in the egg cell.
In polyploid obligate parthenogens like the whiptail lizard, all the offspring are female.[20]
In many hymenopteran insects such as honeybees, female eggs are produced sexually, using sperm from a drone father, while the production of further drones (males) depends on the queen (and occasionally workers) producing unfertilized eggs. This means that females (workers and queens) are always diploid, while males (drones) are always haploid, and produced parthenogenetically.
Facultative
Facultative parthenogenesis is the term for when a female can produce offspring either sexually or via asexual reproduction.[26] Facultative parthenogenesis is extremely rare in nature, with only a few examples of animal taxa capable of facultative parthenogenesis.[26] One of the best-known examples of taxa exhibiting facultative parthenogenesis are mayflies; presumably, this is the default reproductive mode of all species in this insect order.[27] Facultative parthenogenesis is believed to be a response to a lack of a viable male. A female may undergo facultative parthenogenesis if a male is absent from the habitat or if it is unable to produce viable offspring.
Facultative parthenogenesis is often incorrectly used to describe cases of accidental or spontaneous parthenogenesis in normally sexual animals.[28] For example, many cases of accidental parthenogenesis in sharks, some snakes, Komodo dragons and a variety of domesticated birds were widely perpetuated as facultative parthenogenesis.[29] These cases are, however, examples of accidental parthenogenesis, given the frequency of asexually produced eggs and their hatching rates are extremely low, in contrast to true facultative parthenogenesis where the majority of asexually produced eggs hatch.[26][28]  The occurrence of such asexually produced eggs in sexual animals can be explained by a meiotic error, leading to eggs produced via automixis.[28][30]
Obligate
Obligate parthenogenesis is the process in which organisms exclusively reproduce through asexual means.[31] Many species have been shown to transition to obligate parthenogenesis over evolutionary time. Among these species, one of the most well documented transitions to obligate parthenogenesis was found in almost all metazoan taxa, albeit through highly diverse mechanisms. These transitions often occur as a result of inbreeding or mutation within large populations.[32] There are a number of documented species, specifically salamanders and geckos, that rely on obligate parthenogenesis as their major method of reproduction. As such, there are over 80 species of unisex reptiles (mostly lizards but including a single snake species), amphibians and fishes in nature for which males are no longer a part of the reproductive process.[33] A female will produce an ovum with a full set (two sets of genes) provided solely by the mother. Thus, a male is not needed to provide sperm to fertilize the egg. This form of asexual reproduction is thought in some cases to be a serious threat to biodiversity for the subsequent lack of gene variation and potentially decreased fitness of the offspring.[31]
Some invertebrate species that feature (partial) sexual reproduction in their native range are found to reproduce solely by parthenogenesis in areas to which they have been introduced.[34][35] Relying solely on parthenogenetic reproduction has several advantages for an invasive species: it obviates the need for individuals in a very sparse initial population to search for mates, and an exclusively female sex distribution allows a population to multiply and invade more rapidly, potentially up to twice as fast. Examples include several aphid species[34] and the willow sawfly, Nematus oligospilus, which is sexual in its native Holarctic habitat but parthenogenetic where it has been introduced into the Southern Hemisphere.[35]
Natural occurrence
Parthenogenesis is seen to occur naturally in aphids, Daphnia, rotifers, nematodes and some other invertebrates, as well as in many plants. Among vertebrates, strict parthenogenesis is only known to occur in lizards, snakes,[36] birds[37] and sharks,[38] with fish, amphibians and reptiles exhibiting various forms of gynogenesis and hybridogenesis (an incomplete form of parthenogenesis).[39] The first all-female (unisexual) reproduction in vertebrates was described in the fish Poecilia formosa in 1932.[40] Since then at least 50 species of unisexual vertebrate have been described, including at least 20 fish, 25 lizards, a single snake species, frogs, and salamanders.[39] Other, usually sexual, species may occasionally reproduce parthenogenetically and Komodo dragons; the hammerhead and blacktip sharks are recent additions to the known list of spontaneous parthenogenetic vertebrates. As with all types of asexual reproduction, there are both costs (low genetic diversity and therefore susceptibility to adverse mutations that might occur) and benefits (reproduction without the need for a male) associated with parthenogenesis.
Parthenogenesis is distinct from artificial animal cloning, a process where the new organism is necessarily genetically identical to the cell donor. In cloning, the nucleus of a diploid cell from a donor organism is inserted into an enucleated egg cell and the cell is then stimulated to undergo continued mitosis, resulting in an organism that is genetically identical to the donor. Parthenogenesis is different, in that it originates from the genetic material contained within an egg cell and the new organism is not necessarily genetically identical to the parent.
Parthenogenesis may be achieved through an artificial process as described below under the discussion of mammals.
Insects
Parthenogenesis in insects can cover a wide range of mechanisms.[41] The offspring produced by parthenogenesis may be of both sexes, only female (thelytoky, e.g. aphids and some hymenopterans[42]) or only male (arrhenotoky, e.g. most hymenopterans). Both true parthenogenesis and pseudogamy (gynogenesis or sperm-dependent parthenogenesis) are known to occur.[26] The egg cells, depending on the species may be produced without meiosis (apomictically) or by one of the several automictic mechanisms.
A related phenomenon, polyembryony is a process that produces multiple clonal offspring from a single egg cell. This is known in some hymenopteran parasitoids and in Strepsiptera.[41]
In automictic species the offspring can be haploid or diploid. Diploids are produced by doubling or fusion of gametes after meiosis. Fusion is seen in the Phasmatodea, Hemiptera (Aleurodids and Coccidae), Diptera, and some Hymenoptera.[41]
In addition to these forms is hermaphroditism, where both the eggs and sperm are produced by the same individual, but is not a type of parthenogenesis. This is seen in three species of Icerya scale insects.[41]
Parasitic bacteria like Wolbachia have been noted to induce automictic thelytoky in many insect species with haplodiploid systems. They also cause gamete duplication in unfertilized eggs causing them to develop into female offspring.[41]

Among species with the haplo-diploid sex-determination system, such as hymenopterans (ants, bees and wasps) and thysanopterans (thrips), haploid males  are produced from unfertilized eggs. Usually, eggs are laid only by the queen, but the unmated workers may also lay haploid, male eggs either regularly (e.g. stingless bees) or under special circumstances. An example of non-viable parthenogenesis is common among domesticated honey bees. The queen bee is the only fertile female in the hive; if she dies without the possibility of a viable replacement queen, it is not uncommon for the worker bees to lay eggs. This is a result of the lack of the queen's pheromones and the pheromones secreted by uncapped brood, which normally suppress ovarian development in workers. Worker bees are unable to mate, and the unfertilized eggs produce only drones (males), which can mate only with a queen. Thus, in a relatively short period, all the worker bees die off, and the new drones follow if they have not been able to mate before the collapse of the colony. This behavior is believed to have evolved to allow a doomed colony to produce drones which may mate with a virgin queen and thus preserve the colony's genetic progeny.
A few ants and bees are capable of producing diploid female offspring parthenogenetically. These include a honey bee subspecies from South Africa, Apis mellifera capensis, where  workers are capable of producing diploid eggs parthenogenetically, and replacing the queen if she dies; other examples include some species of small carpenter bee, (genus Ceratina). Many parasitic wasps are known to be parthenogenetic, sometimes due to infections by Wolbachia.
The workers in five[23] ant species and the queens in some ants are known to reproduce by parthenogenesis. In Cataglyphis cursor, a European formicine ant, the queens and workers can produce new queens by parthenogenesis. The workers are produced sexually.[23]
In Central and South American electric ants, Wasmannia auropunctata, queens produce more queens through automictic parthenogenesis with central fusion. Sterile workers usually are produced from eggs fertilized by males. In some of the eggs fertilized by males, however, the fertilization can cause the female genetic material to be ablated from the zygote. In this way, males pass on only their genes to become fertile male offspring. This is the first recognized example of an animal species where both females and males can reproduce clonally resulting in a complete separation of male and female gene pools.[43] As a consequence, the males will only have fathers and the queens only mothers, while the sterile workers are the only ones with both parents of both genders.
These ants get both the benefits of both asexual and sexual reproduction[23][43]—the daughters who can reproduce (the queens) have all of the mother's genes, while the sterile workers whose physical strength and disease resistance are important are produced sexually.
Other examples of insect parthenogenesis can be found in gall-forming aphids (e.g., Pemphigus betae), where females reproduce parthenogenetically during the gall-forming phase of their life cycle and in grass thrips. In the grass thrips genus Aptinothrips there have been, despite the very limited number of species in the genus, several transitions to asexuality.[44]
Crustaceans
Crustacean reproduction varies both across and within species. The water flea Daphnia pulex alternates between sexual and parthenogenetic reproduction.[45]  Among the better-known large decapod crustaceans, some crayfish reproduce by parthenogensis. "Marmorkrebs" are parthenogenetic crayfish that were discovered in the pet trade in the 1990s.[46]  Offspring are genetically identical to the parent, indicating it reproduces by apomixis, i.e. parthenogenesis in which the eggs did not undergo meiosis.[47] Spinycheek crayfish (Orconectes limosus) can reproduce both sexually and by parthenogenesis.[48] The Louisiana red swamp crayfish (Procambarus clarkii), which normally reproduces sexually, has also been suggested to reproduce by parthenogenesis,[49] although no individuals of this species have been reared this way in the lab. Artemia parthenogenetica is a species or series of populations of parthenogenetic brine shrimps.[50]
Velvet worms
No males of Epiperipatus imthurni have been found, and specimens from Trinidad were shown to reproduce parthenogenetically. This species is the only known velvet worm to reproduce via parthenogenesis.[51]
Spiders
At least two species of spiders in the family Oonopidae (goblin spiders), Heteroonops spinimanus and Triaeris stenaspis, are thought to be parthenogenetic, as no males have ever been collected. Parthenogenetic reproduction has been demonstrated in the laboratory for T.stenaspis.[52]
Rotifers
In bdelloid rotifers, females reproduce exclusively by parthenogenesis (obligate parthenogenesis),[53]  while in monogonont rotifers, females can alternate between sexual and asexual reproduction (cyclical parthenogenesis). At least in one normally cyclical parthenogenetic species obligate parthenogenesis can be inherited: a recessive allele leads to loss of sexual reproduction in homozygous offspring.[54]
Flatworms
At least two species in the genus Dugesia, flatworms in the Turbellaria sub-division of the phylum Platyhelminthes, include polyploid individuals that reproduce by parthenogenesis.[55] This type of parthenogenesis requires mating, but the sperm does not contribute to the genetics of the offspring (the parthenogenesis is pseudogamous, alternatively referred to as gynogenetic). A complex cycle of matings between diploid sexual and polyploid parthenogenetic individuals produces new parthenogenetic lines.
Snails
Several species of parthenogenetic gastropods have been studied, especially with respect to their status as invasive species. Such species include the New Zealand mud snail (Potamopyrgus antipodarum),[56] the red-rimmed melania (Melanoides tuberculata),[57] and the Quilted melania (Tarebia granifera).[58]
Squamata

Most reptiles of the squamatan order (lizards and snakes) reproduce sexually, but parthenogenesis has been observed to occur naturally in certain species of whiptails, some geckos, rock lizards,[5][59][60] Komodo dragons and snakes.[61] Some of these like the mourning gecko Lepidodactylus lugubris, Indo-Pacific house gecko Hemidactylus garnotii, the hybrid whiptails Cnemidophorus, Caucasian rock lizards Darevskia, and the brahminy blindsnake, Indotyphlops braminus are unisexual and obligately parthenogenetic. Other reptiles, such as the Komodo dragon, other monitor lizards,[62] and some species of boas,[8][24][63] pythons,[22][64] filesnakes,[65][66] gartersnakes[67] and rattlesnakes[68][69] were previously considered as cases of facultative parthenogenesis, but are in fact cases of accidental parthenogenesis.[28]
In 2012, facultative parthenogenesis was reported in wild vertebrates for the first time by US researchers amongst captured pregnant copperhead and cottonmouth female pit-vipers.[70] The Komodo dragon, which normally reproduces sexually, has also been found able to reproduce asexually by parthenogenesis.[71] A case has been documented of a Komodo dragon reproducing via sexual reproduction after a known parthenogenetic event,[72] highlighting that these cases of parthenogenesis are reproductive accidents, rather than adaptive, facultative parthenogenesis.[28]
Some reptile species use a ZW chromosome system, which produces either males (ZZ) or females (ZW). Until 2010, it was thought that the ZW chromosome system used by reptiles was incapable of producing viable WW offspring, but a (ZW) female boa constrictor was discovered to have produced viable female offspring with WW chromosomes.[73]
Parthenogenesis has been studied extensively in the New Mexico whiptail in the genus Aspidoscelis of which 15 species reproduce exclusively by parthenogenesis. These lizards live in the dry and sometimes harsh climate of the southwestern United States and northern Mexico. All these asexual species appear to have arisen through the hybridization of two or three of the sexual species in the genus leading to polyploid individuals. The mechanism by which the mixing of chromosomes from two or three species can lead to parthenogenetic reproduction is unknown. Recently, a hybrid parthenogenetic whiptail lizard was bred in the laboratory from a cross between an asexual and a sexual whiptail.[74] Because multiple hybridization events can occur, individual parthenogenetic whiptail species can consist of multiple independent asexual lineages. Within lineages, there is very little genetic diversity, but different lineages may have quite different genotypes.
An interesting aspect to reproduction in these asexual lizards is that mating behaviors are still seen, although the populations are all female. One female plays the role played by the male in closely related species, and mounts the female that is about to lay eggs. This behaviour is due to the hormonal cycles of the females, which cause them to behave like males shortly after laying eggs, when levels of progesterone are high, and to take the female role in mating before laying eggs, when estrogen dominates. Lizards who act out the courtship ritual have greater fecundity than those kept in isolation, due to the increase in hormones that accompanies the mounting. So, although the populations lack males, they still require sexual behavioral stimuli for maximum reproductive success.[75]
Some lizard parthenogens show a pattern of geographic parthenogenesis, occupying high mountain areas where their ancestral forms have an inferior competition ability.[76] In Caucasian rock lizards of genus Darevskia, which have six parthenogenetic forms of hybrid origin[59][60][77] hybrid parthenogenetic form D. "dahli" has a broader niche than either of its bisexual ancestors and its expansion throughout the Central Lesser Caucasus caused decline of the ranges of both its maternal and paternal species.[78]
Amphibians

Sharks
Parthenogenesis in sharks has been confirmed in at least three species, the bonnethead,[38] the blacktip shark,[79] and the zebra shark,[80]  and reported in others.
A bonnethead, a type of small hammerhead shark, was found to have produced a pup, born live on 14 December 2001 at Henry Doorly Zoo in Nebraska, in a tank containing three female hammerheads, but no males. The pup was thought to have been conceived through parthenogenesis. The shark pup was apparently killed by a stingray within days of birth.[81] The investigation of the birth was conducted by the research team from Queen's University Belfast, Southeastern University in Florida, and Henry Doorly Zoo itself, and it was concluded after DNA testing that the reproduction was parthenogenetic. The testing showed the female pup's DNA matched only one female who lived in the tank, and that no male DNA was present in the pup. The pup was not a twin or clone of her mother, but rather, contained only half of her mother's DNA ("automictic parthenogenesis"). This type of reproduction had been seen before in bony fish, but never in cartilaginous fish such as sharks, until this documentation.
In the same year, a female Atlantic blacktip shark in Virginia reproduced via parthenogenesis.[82] On 10 October 2008 scientists confirmed the second case of a virgin birth in a shark. The Journal of Fish Biology reported a study in which scientists said DNA testing proved that a pup carried by a female Atlantic blacktip shark in the Virginia Aquarium & Marine Science Center contained no genetic material from a male.[79]
In 2002, two white-spotted bamboo sharks were born at the Belle Isle Aquarium in Detroit. They hatched 15 weeks after being laid. The births baffled experts as the mother shared an aquarium with only one other shark, which was female. The female bamboo sharks had laid eggs in the past. This is not unexpected, as many animals will lay eggs even if there is not a male to fertilize them. Normally, the eggs are assumed to be inviable and are discarded. This batch of eggs was left undisturbed by the curator as he had heard about the previous birth in 2001 in Nebraska and wanted to observe whether they would hatch. Other possibilities had been considered for the birth of the Detroit bamboo sharks including thoughts that the sharks had been fertilized by a male and stored the sperm for a period of time, as well as the possibility that the Belle Isle bamboo shark is a hermaphrodite, harboring both male and female sex organs, and capable of fertilizing its own eggs, but that is not confirmed.[83]
In 2008, a Hungarian aquarium had another case of parthenogenesis after its lone female shark produced a pup without ever having come into contact with a male shark.
The repercussions of parthenogenesis in sharks, which fails to increase the genetic diversity of the offspring, is a matter of concern for shark experts, taking into consideration conservation management strategies for this species, particularly in areas where there may be a shortage of males due to fishing or environmental pressures. Although parthenogenesis may help females who cannot find mates, it does reduce genetic diversity.
In 2011, recurring shark parthenogenesis over several years was demonstrated in a captive zebra shark, a type of carpet shark.[80][84] DNA genotyping demonstrated that individual zebra sharks can switch from sexual to parthenogenetic reproduction.[85]
Birds
Parthenogenesis in birds is known mainly from studies of domesticated turkeys and chickens, although it has also been noted in the domestic pigeon.[37] In most cases the egg fails to develop normally or completely to hatching.[37][86] The first description of parthenogenetic development in a passerine was demonstrated in captive zebra finches, although the dividing cells exhibited irregular nuclei and the eggs did not hatch.[37]
Parthenogenesis in turkeys appears to result from a conversion of haploid cells to diploid;[86] most embryos produced in this way die early in development. Rarely, viable birds result from this process, and the rate at which this occurs in turkeys can be increased by selective breeding,[87] however male turkeys produced from parthenogenesis exhibit smaller testes and reduced fertility.[88]
Mammals
There are no known cases of naturally occurring mammalian parthenogenesis in the wild. Parthenogenetic progeny of mammals would have two X chromosomes, and would therefore be female.
In 1936, Gregory Goodwin Pincus reported successfully inducing parthenogenesis in a rabbit.[89]
Helen Spurway, a geneticist specializing in the reproductive biology of the guppy, Lebistes reticulatus, claimed, in 1955, that parthenogenesis, which occurs in the guppy in nature, may also occur (though very rarely) in the human species, leading to so-called "virgin births."  This created some sensation among her colleagues and the lay public alike.[90]
In April 2004, scientists at Tokyo University of Agriculture used parthenogenesis successfully to create a fatherless mouse. Using gene targeting, they were able to manipulate two imprinted loci H19/IGF2 and DLK1/MEG3 to produce bi-maternal mice at high frequency[91]  and subsequently show that fatherless mice have enhanced longevity.[92]
Induced parthenogenesis in mice and monkeys often results in abnormal development. This is because mammals have imprinted genetic regions, where either the maternal or the paternal chromosome is inactivated in the offspring in order for development to proceed normally. A mammal created by parthenogenesis would have double doses of maternally imprinted genes and lack paternally imprinted genes, leading to developmental abnormalities. It has been suggested[93]  that defects in placental folding or interdigitation are one cause of swine parthenote abortive development. As a consequence, research on human parthenogenesis is focused on the production of embryonic stem cells for use in medical treatment, not as a reproductive strategy.
Use of an electrical or chemical stimulus can produce the beginning of the process of parthenogenesis in the asexual development of viable offspring.[94]

During oocyte development, high metaphase promoting factor (MPF) activity causes mammalian oocytes to arrest at the metaphase II stage until fertilization by a sperm. The fertilization event causes intracellular calcium oscillations, and targeted degradation of cyclin B, a regulatory subunit of MPF, thus permitting the MII-arrested oocyte to proceed through meiosis.
To initiate parthenogenesis of swine oocytes, various methods exist to induce an artificial activation that mimics sperm entry, such as calcium ionophore treatment, microinjection of calcium ions, or electrical stimulation. Treatment with cycloheximide, a non-specific protein synthesis inhibitor, enhances parthenote development in swine presumably by continual inhibition of MPF/cyclin B.[95] As meiosis proceeds, extrusion of the second polar is blocked by exposure to cytochalasin B. This treatment results in a diploid (2 maternal genomes) parthenote[93] Parthenotes can be surgically transferred to a recipient oviduct for further development, but will succumb to developmental failure after ≈30 days of gestation. The swine parthenote placentae often appears hypo-vascular: see free image (Figure 1) in linked reference.[93]
Humans
On June 26, 2007, International Stem Cell Corporation (ISCC), a California-based stem cell research company, announced that their lead scientist, Dr. Elena Revazova, and her research team were the first to intentionally create human stem cells from unfertilized human eggs using parthenogenesis. The process may offer a way for creating stem cells that are genetically matched to a particular female for the treatment of degenerative diseases that might affect her. In December 2007, Dr. Revazova and ISCC published an article[96] illustrating a breakthrough in the use of parthenogenesis to produce human stem cells that are homozygous in the HLA region of DNA. These stem cells are called HLA homozygous parthenogenetic human stem cells (hpSC-Hhom) and have unique characteristics that would allow derivatives of these cells to be implanted into millions of people without immune rejection.[97] With proper selection of oocyte donors according to HLA haplotype, it is possible to generate a bank of cell lines whose tissue derivatives, collectively, could be MHC-matched with a significant number of individuals within the human population.
On August 2, 2007, after a much independent investigation, it was revealed that discredited South Korean scientist Hwang Woo-Suk unknowingly produced the first human embryos resulting from parthenogenesis. Initially, Hwang claimed he and his team had extracted stem cells from cloned human embryos, a result later found to be fabricated. Further examination of the chromosomes of these cells show indicators of parthenogenesis in those extracted stem cells, similar to those found in the mice created by Tokyo scientists in 2004. Although Hwang deceived the world about being the first to create artificially cloned human embryos, he did contribute a major breakthrough to stem cell research by creating human embryos using parthenogenesis.[98] The truth was discovered in 2007, long after the embryos were created by him and his team in February 2004. This made Hwang the first, unknowingly, to successfully perform the process of parthenogenesis to create a human embryon and, ultimately, a human parthenogenetic stem cell line.
Oomycetes
Apomixis can apparently occur in Phytophthora,[99] an oomycete. Oospores from an experimental cross were germinated, and some of the progeny were genetically identical to one or other parent, implying that meiosis did not occur and the oospores developed by parthenogenesis.
Similar phenomena
Gynogenesis

A form of asexual reproduction related to parthenogenesis is gynogenesis. Here, offspring are produced by the same mechanism as in parthenogenesis, but with the requirement that the egg merely be stimulated by the presence of sperm in order to develop. However, the sperm cell does not contribute any genetic material to the offspring. Since gynogenetic species are all female, activation of their eggs requires mating with males of a closely related species for the needed stimulus. Some salamanders of the genus Ambystoma are gynogenetic and appear to have been so for over a million years. It is believed that the success of those salamanders may be due to rare fertilization of eggs by males, introducing new material to the gene pool, which may result from perhaps only one mating out of a million. In addition, the amazon molly is known to reproduce by gynogenesis.[100]
Hybridogenesis

Hybridogenesis is a mode of reproduction of hybrids. Hybridogenetic hybrids (for example AB genome), usually females, during gametogenesis exclude one of parental genomes (A) and produce gametes with unrecombined[101] genome of second parental species (B), instead of containing mixed recombined parental genomes. First genome (A) is restored by fertilization of these gametes with gametes from the first species (AA, sexual host,[101] usually male).[101][102][103]
So hybridogenesis is not completely asexual, but instead hemiclonal: half of genome is passed to the next generation clonally, unrecombined, intact (B), other half sexually, recombined (A).[101][104]
This process continues, so that each generation is half (or hemi-) clonal on the mother's side and has half new genetic material from the father's side.
This form of reproduction is seen in some live-bearing fish of the genus Poeciliopsis[102][105] as well as in some of the Pelophylax spp. ("green frogs" or "waterfrogs"):
P. kl. esculentus (edible frog): P. lessonae × P. ridibundus,[101][106][107]
P. kl. grafi (Graf's hybrid frog): P. perezi × P. ridibundus[101]
P. kl. hispanicus (Italian edible frog) – unknown origin: P. bergeri × P. ridibundus or P. kl. esculentus[101]
and perhaps in P. demarchii.

Other examples where hybridogenesis is at least one of modes of reproduction include i.e.
Iberian minnow Tropidophoxinellus alburnoides (Squalius pyrenaicus × hypothetical ancestor related with Anaecypris hispanica)[109]
spined loaches Cobitis hankugensis × C. longicorpus[110]
Bacillus stick insects B. rossius × Bacillus grandii benazzii[111]
See also
Charles Bonnet – conducted experiments that established what is now termed parthenogenesis in aphids
Jacques Loeb – caused the eggs of sea urchins to begin embryonic development without sperm
Miraculous births
Parthenocarpy – plants with seedless fruit

Further reading
Dawley, Robert M. & Bogart, James P. (1989). Evolution and Ecology of Unisexual Vertebrates. Albany, New York: New York State Museum. ISBN1-55557-179-4.

Futuyma, Douglas J. & Slatkin, Montgomery. (1983). Coevolution. Sunderland, Mass: Sinauer Associates. ISBN0-87893-228-3.

Maynard Smith, John. (1978). The Evolution of Sex. Cambridge: Cambridge University Press. ISBN0-521-29302-2.
Michod, Richard E. & Levin, Bruce R. (1988). The Evolution of Sex. Sunderland, Mass: Sinauer Associates. ISBN0-87893-459-6.

Stearns, Stephan C. (1988). The Evolution of Sex and Its Consequences (Experientia Supplementum, Vol. 55). Boston: Birkhauser. ISBN0-8176-1807-4.

 from Oregon State University


This is a list of Latin American and Caribbean countries by gross domestic product at purchasing power parity in international dollars according to the International Monetary Fund's estimates in the October 2018 World Economic Outlook database.
The Latin American countries Brazil, Mexico, and Argentina accounted for over two-thirds of the region's total gross domestic product (GDP) at purchasing power parity (PPP) in 2019, while Caribbean and North Atlantic nations represented just over 1% of the region's total GDP (PPP). The Bahamas had the region's highest GDP (PPP) per capita, whereas Haiti had the lowest.
Cuba is not included in the list due to lack of economic data. Puerto Rico is not listed since it is a U.S. territory.

See also

Community of Latin American and Caribbean States
List of Latin American and Caribbean countries by GDP (nominal)
List of Latin American and Caribbean countries by GDP growth

List of Latin American and Caribbean countries by GDP (PPP)

What is the poorest country in Latin America?


As of January 1, 2011, The Church of Jesus Christ of Latter-day Saints reported 38,257 members in 14 stakes and 3 districts, 173 congregations (89 wards[1] and 84 branches[1]), three missions, and two temples in Germany.[2]  In 2017, according to the Church's Mormon Newsroom web site, there were 40,011 members in 183 congregations.
History

The first German to be converted to the LDS Church was an immigrant to the USA named Jakob Zundel in 1836.[3]
Although one British Mormon convert had briefly worked in Germany, the first official of the church to arrive in Germany was Orson Hyde on 27 June 1841 as part of his journey to Palestine. He was delayed in Frankfurt by a visa problem and began to learn German. When he returned to Germany on his trip back from Palestine, he spent 30 January to August 1842 in Regensburg and wrote "Ein Ruf aus der Wüste" ("A Cry out of the Wilderness") whilst he was there. It was published in Frankfurt and was the first LDS Church publication in the German language.[3]
The first converts were baptized in Germany in 1851. Brigham Young sent Daniel Carn to establish the first German mission in 1852 which he did in Hamburg. Carn also oversaw the publication of a German language version of the Book of Mormon which was published in Hamburg 25 May 1852. He was eventually banished from Hamburg, then a sovereign state, due to his attempts to convert Germans to Mormonism but he continued to prostelyse Germans in the then Danish territory of Schleswig-Holstein.[3]
Most early converts emigrated to the United States and, by 1854, the short-lived Hamburg branch was dissolved.  Church involvement in Germany resumed in 1860, but was limited due to persecution which continued until World War I. After World War I, the government became more tolerant of religious freedom, and the church received substantial growth.
In the first half of the 20th century Germany had more converts to Mormonism than any other non-English speaking country.[4] By 1925 there were 6,125 members in the German-Austrian Mission, and 5,305 members in the Swiss-German Mission. The first German LDS meeting house was built in 1929 in Selbongen, East Prussia (now Zełwągi in Poland).[5][6]
13,402 Mormons lived in the West German (including Austria as of November 1938) and East German Missions in 1939; at least 996 members were killed during World War II, including more than 400 adult men, about 10% of priesthood holders.[7] Under the Nazi Government of 1933 - 1945, no Mormon congregation was stopped from worshipping and few individual Mormons were persecuted (and only for transgressions that any German of the time would have been punished for).[8] Gestapo agents silently attended services, likely investigating neighbors' complaints of seditious activities, but no punishment came to the church. Adhering to the 12th Article of Faith, an estimated 3-5% of adult male members joined the Nazi Party—required of state employees—and speakers avoided criticizing the government or, after the German declaration of war against the United States, emphasizing the church's relationship with that country. The government ordered the church to avoid preaching about "Jewish" topics like "Zion" and "Israel", so leaders told members to not sing hymns with such words.[7] American Mormon missionaries' views of the government during the 1930s varied. While praising Adolf Hitler's oratory skill and approving of his unifying a politically divided country, they saw arrests of dissidents, enforcement of Nazi eugenics, and widespread fear of the regime. The Nuremberg Laws increased access to and interest in genealogical records, and some saw the monthly eintopf as similar to Fast Sunday, but mandatory Hitler Youth membership ended most Mormon auxiliary organizations for young people.[9]
Following World War II, members of the church in Germany found themselves divided among two nations. Members continued to maintain contact with the church in the west. In the fall of 1961 three stakes were created in Berlin (Germany's first), Stuttgart and Hamburg. In 1982, the Freiberg German Democratic Republic Stake was created. On June 19, 1985, the Freiberg GDR Temple was dedicated. It is the only temple constructed in a communist bloc country. In 1987, the Frankfurt, West Germany Temple was dedicated.[2][10]
Missions
Germany Berlin Mission
Germany Frankfurt Mission
Alpine German-speaking Mission (also covers Austria and parts of Switzerland)(does not exclude areas of southern Germany)
Temples

See also

Religion in Germany

 at L. Tom Perry Special Collections, Harold B. Lee Library, Brigham Young University

 - Official Site
 - Visitors Site

The Church of Jesus Christ of Latter-day Saints in Germany


Part of the Politics seriesBasic forms of government
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A right-wing dictatorship (sometimes also referred to as a rightist dictatorship) is an authoritarian (or sometimes totalitarian) regime whose policy could be called right-wing. There are various definitions of the term "rightist". The broadest one includes all dictatorships that do not consider themselves communist. Those are usually pro-capitalist in economic matters and conservative in ideological ones. The term fascist dictatorship is sometimes used interchangeably with the right-wing one. It is commonly accepted that Nazi Germany and the Kingdom of Italy were ruled by fascist governments at some points of their history, but how it refers to other right-wing regimes is a question of further debate. The Estado Novo in Portugal was a right-wing dictatorship which was corporatist in nature. Most South American dictatorships during the second half of the 20th century were right-wing: Pinochet, the Brazilian Military Government, etc. There have also been a number of military dictatorships installed by anti-communists which were generally conservative and rightist.
Military government
In the most common Western view, the perfect example of a right-wing dictatorship is any of those that once ruled in South America. Those regimes were predominantly military juntas and most of them collapsed in the 1980s. Communist countries, which were very cautious about not revealing their authoritarian methods of rule to the public, were usually led by civilian governments and officers taking power were not much welcomed there. Few exceptions include the Burmese Way to Socialism (Burma, 1966–1988), the Military Council of National Salvation (People's Republic of Poland, 1981–1983) or the North Korean regime's evolution throughout the rule of Kim Il-sung.
Religion and the government
Most right-wing regimes kept strong ties with local Churches (usually the Roman Catholic ones since most of those regimes happened in Catholic countries). This policy of a strong Church-state alliance is usually referred to as Austrofascism. The most pro-Catholic dictatorships were Portugal (1933–1974) and the Federal State of Austria (1934–1938). Non-Christian dictatorships include those in the Muslim world, the most famous being Iran since the revolution of 1979. There are several other examples of theocratic (and therefore right-wing) regimes in the region, like Somalia or Afghanistan under the Taliban. While it is unclear whether a monarchy could be called a dictatorship, theocratic absolute monarchies of Saudi Arabia or Vatican City share many similarities with the regimes mentioned above. Many of those are/were led by spiritual leaders and examples include the Slovak Republic under the Reverend Josef Tiso or Iran under the Ayatollahs Khomeini (1979–1989) and Khamenei (1989–present). Some right-wing dictatorships, like the Nazi Germany, were even openly hostile to certain religions.[1]
Europe
In some post-communist countries, people involved in the previous regime took power in the years following the end of the Cold War. The most famous example of a European dictatorship of that sort is Belarus under Alexander Lukashenko. Belarus and Azerbaijan were both considered "authoritarian regimes" by the Democracy Index as of 2008. Freedom House marks them "not-free states" together with Russia, so whether Russia is a dictatorship or just a seriously flawed democracy is a question of further debate. All three governments are non-communist and especially in Russia sometimes stay in total opposition to the communist parties, therefore they could be called right-wing as well.
List of European right-wing dictatorships
Considering a right-wing dictatorship any non-democratic (non-parliamentary or pseudo-parliamentary political system, suppression of basic freedoms such as association and speech, prosecution of opponents, etc.) regime that is not communist, the list of European examples would be as following:
Current

Former

 Semi-authoritarian regime.
 Semi-authoritarian regime in the years 1938–1940.
 Semi-authoritarian regime.
Asia
List of Asian former right-wing dictatorships

See also
Films depicting Latin American military dictatorships
List of political leaders who held active military ranks in office
Military dictatorship
Military government
Police state

What is the most violent dictatorship in history?


This is a list of Latin American countries and dependent territories by population, which is sorted by the 2015 mid-year normalized demographic projections.
Table

Notes

See also
List of countries by population
List of African countries by population
List of countries in the Americas by population
List of Arab countries by population
List of Asian countries by population
List of Caribbean island countries by population
List of Eurasian countries by population
List of European countries by population
List of European Union member states by population
List of member states of the Commonwealth of Nations by population
List of Middle East countries by population
List of North American countries by population
List of Oceanian countries by population
List of South American countries by population
List of countries by past and future population
List of countries by past population (United Nations)
List of countries by population in 2000
List of countries by population in 2010

List of Latin American countries by population

What is the largest country in Latin America?


The economy of Hyderabad, the capital of Telangana, India, is based on traditional manufacturing, the knowledge sector, and tourism.[1] Starting in the 1990s, the economic pattern of the city changed from a primarily service hub to a more diversified economy,[2] but the service industry remains a major contributor.[3] As of 2006, the largest employers of Hyderabad are the governments of Andhra Pradesh and India, with 113,000 and 85,000 employees, respectively.[4]

Since its inception in 1591, Hyderabad has been a global trade center in multiple areas, including its status as the world's only diamond market. City-based handicrafts were sold in the Middle East and Western countries. During Nizam's rule in the 1930s, industrial growth started with the establishment of a diversified industrial zone, which grew in parallel with traditional manufacturing.
In the 1930s and 1940s, city-based industries started importing technology from the western world for industrial manufacturing. With the introduction of the railways, the city became well connected with the port cities of Bombay (Mumbai), Madras (now Chennai), Calcutta (now Kolkata) and Karachi (now in Pakistan).[5][6] During the 1950s and 1960s, most of the Indian premier public enterprises—BHEL, NMDC, HMT, BEL, IDPL, ECIL, DRDO and HAL—were established in Hyderabad, changing the economical pattern of the city from a traditional manufacturing to a cosmopolitan industrial service sector.[7]
Hyderabad is the capital of Telangana, and the city is the largest contributor to the state's GDP (Gross domestic product) and state tax.[8] In 2011, Hyderabad generated revenues of ₹700,000 million (US$10billion) and contributed a third of the state's tax revenue.[9] In 2008, the GDP (PPP) was US$60bn, placing the city fourth in India and 93rd in the world.[10][3] Hyderabad and its suburbs house the highest number of special economic zones among India's cities.[11]
In the 1970s, the pharmaceutical and electronic industries were established in the city because of its strategic location in south-central India, for which it is known as the gateway to south-central India.[7][12][13] Since the 1990s, the economic patterns of the city have changed it from a primarily service hub to a more diversified spectrum,[2] with the growth of IT enterprises, and biotech, insurance, and financial institutions, and a strong employment base in ancillary activities such as trade and commerce, transport, storage, communication, real-estate and retail, which employ three times more people than the IT industries.[6] The service industry in this arena remains a dominant, with 90% of the workforce.[13][14] As of 2005, out of every 1000 people of working age, 770 males and 190 females are employed.[15]
Tourism

In March 2012, The Indian Union Tourism Ministry declared Hyderabad as the first ever "Best Heritage City of India".[16]
Hyderabad was the World's third best city to visit in 2013, according to Lonely Planet.  In 2011 the city was ranked nineteenth in the world by The New York Times in The list of 41 Places to Go in 2011, the only Indian City in the list.[17] Hyderabad is known as the City of Pearls, due to the presence of pearls trading industry—until the 18th century the city was the only global trade center of large diamonds.[18][19] Many traditional and historical bazaars are located in the city.[20][21] The Laad Bazaar and nearby markets have shops that sell pearls, diamonds and other traditional ware and cultural antiques.[20]
Biopharmaceutical industry

Hyderabad emerged as a pharmaceutical and biotechnology hub and is known as India's pharmaceutical capital and "Genome Valley of India".[22][23] In 2008–2009, Hyderabad's bio-pharmaceuticals exports reached US$3.1billion.[24] The south—central location of Hyderabad, and the incorporation of Indian Drugs and Pharmaceuticals Limited (IDPL) in 1961 proved to be a foundation of pharmaceutical industry in Hyderabad, later in the 1990s the expansion in the industry took place with the formation of Indian Institute of Chemical Technology, National Institute of Pharmaceutical Education and Research, the Centre for Cellular and Molecular Biology and National Institute of Nutrition along with the regional institutions making Hyderabad the hub of pharmaceutical and biotechnology in India.[25]
The establishment of the public sector in biotechnology and the Genome Valley, 'Fab City' and the 'Nano Technology park' established significant infrastructure in bio-technology. These attributes attracted regional companies and MNCs to set up offices, warehouses, research and development centres in the city.[26] In 2012, the city hosted the Convention on Biological Diversity (2012 COP 11).[27]
The commercial market structure of Hyderabad is defined into 4 sectors—The Central Business Districts (CBD), the sub-central business centres, the neighbourhood business centres and local business centres.[28] Since 2007, The retail industry in Hyderabad is on the rise,[29] and several central business districts are spread across the city.[30][31]

Retail and real estate
The World Bank Group ranked the city as the second best Indian city for doing business in 2009.[32] In 2010, the economic analysis group GaWC ranked Hyderabad in its third tier (Gamma+ World City) of cities by importance.[33] In 2011, DTZ ranked Hyderabad as world's third most affordable office location,[34] while Business Today ranked Hyderabad as the fourth best city to live in India.[35] Hyderabad witnessed a high growth in the real estate business, making it among the top five concentrated cities for housing in India.[36] In 2007–08, the city's prime residential areas of Banjara Hills and Jubilee Hills reached the highest growth percentage in India. The Economic Times evaluated Banjara Hills to be worth US$20.7billion.[37]
Information technology

Hyderabad is among the global centres of information technology for which it is known as Cyberabad (Cyber City).[38][39] The city's IT sector includes the IT-enabled services, business process outsourcing, entertainment industries,[39] and Financial services.[40] During 2008–09, Hyderabad's IT exports reached US$ 4.7billion,[24] and 22% of the NASSCOM's total membership is from Hyderabad.[11] The development of a township with related technological infrastructure called HITEC City prompted global and particularly US—based companies to establish their operations in Hyderabad.[38] The Deloitte, Franklin Templeton Investments, GE Capital, Accenture, HSBC, Bank of America, ABN Amro, S&P Capital IQ, Ernst & Young, KPMG, Capgemini, Genpact are some of the financial services companies with offices in the city.[40]
The major multinational IT firms located in Hyderabad are Microsoft (the largest R&D campus outside the US), Google, Thoughtworks, CA Technologies, Amazon.com, IBM, Motorola, Samsung, Agilent, Automatic Data Processing, Oracle Corporation, Yahoo!, Dell, Texas Instruments, Hewlett-Packard, Virtusa, Kewill, Facebook and others.[40][41] The major Indian firms with development centres in the city are Tech Mahindra, Infosys, Wipro, Cognizant, Tata Consultancy Services, Polaris, and others.[40] The main areas where such IT and ITeS campuses have been set up are Madhapur, Gachibowli, Kondapur and Uppal.[42] Apple announced the opening of a new office in Hyderabad that will focus on development of its Maps product.[43]
Hyderabad was conferred the title the 'software training capital of India-and rightly so. The role played by the city of Hyderabad to train the number of software professionals, who get sent to different parts of the country and around the world, have enabled the boom in the software industry in India that we see today. Since 1990, Hyderabad has seen an upward surge in the number of multinationals and tertiary sector services, making the city a tough one to beat, in terms of IT advancement. Many domestic firms and multinational companies have established their headquarters in this city and have contributed to the steady growth of the tertiary sector in Andhra Pradesh. The boom in the software industry has even led to the growth of Hyderabad's very own IT parks such as HITEC City and Mindspace Cyberabad. If you want to know more about the thriving IT industry in Hyderabad, read on.
Types Of IT Companies
There are three main types of IT companies in Hyderabad. They are IT companies, ITES companies and Computer Hardware companies. The IT companies handle the various software development programs for the multiple industrial sectors in Hyderabad and the rest of the world. In this category, Hyderabad has a colossal number of domestic and international companies to its credit. This segment of the Hyderabad IT industry earns the maximum export revenue for the Indian government. The ITES companies (IT enabled services) deal largely with BPO companies and support services. This segment also forms a major chunk of the IT industry in Hyderabad. The third and the final segment of IT companies are the computer hardware companies. Several brands of computer hardware companies provide support to multinational companies and domestic ones and are in the bid to expand their network to the rest of the country.
Major IT Companies
Some of the major IT companies in Hyderabad are:
Google Inc.
IBM
Microsoft
Infosys
SAP
Tata Consultancy Services
Amazon Inc.
Hewlett-Packard
Sony
Toshiba
Nokia
Ericson
Texas Instruments
Ogilvy And Mather
Dell
Deloitte
Governmental Support
The government has done much to support the development and the growth of the IT industry in Hyderabad. Many multinational companies receive invitations from the Government of Andhra Pradesh and India to come and set up their headquarters/ base for operations in the country. The government also provides support in the land allocations aspect in the event of building an IT park or establishing a single office. Many companies benefited from the building of the first IT Park in the city, HITEC City. Apart from land allocation aid, the government also helps domestic and international companies avail state-of-the-art facilities with modern offices and international styled architecture.
Hyderabad provides an encouraging environment, getting various IT companies to establish their bases, grow and network with one another across the world. One of the fastest growing economies owing to the IT sector, Hyderabad is home to some of the biggest multinational companies, BPO's and lavish IT parks.
The future of the IT industry in Hyderabad promises growth and success stories along with expansion and further technological development. It is said that Hyderabad is also known as the second Silicon Valley of India, after Bangalore. Widespread investments, digital infrastructures and training centers are aiding the growth of the number of IT parks around the city. The city has also been selected to house India's first silicon developing facility known as Fab City making Hyderabad truly India's high-tech city.
Hyderabad has several congenial factors to further fuel growth in IT sector. It has earned global reputation as a cosmopolitan city known for its assimilative cultural ethos. Firm foundation, futuristic policy, strategic advantages associated with the city etc., would significantly contribute to the enhancement and enrichment of the IT landscape of Hyderabad
The information and communication technology (ICT) has transformed the image of Hyderabad from a manufacturing and research hub to that of a global services destination. The city could attract many Indian and foreign IT majors thus integrating itself firmly with Silicon Valley . The National Association of Software and Services Companies (NASSCOM) estimates that for every one direct employment, this sector provides indirect employment to four people.
The consumption patterns of middle class youth employed in this sector opens up new avenues of economic opportunities in the city. The brand image this sector brings to the city outweighs the low level of employment elasticity in this knowledge economy, when compared to the bricks and mortar economy. However, it is wrong to assume that IT and IT-Enabled Sectors (ITES) sectors provide employment to elite only. Though the urban and English educated have an edge over others, many lower-middle-class families coming from small towns of Telangana did find their lives significantly impacted by the employment in this sector.
As the Chief Minister, K.Chandrashekar Rao, said while unveiling the new ICT policy, IT has the potential to usher in rapid progress.  Hyderabad is  already a magnet which attracts the best in the world. The new ICT policy intends to embellish it even further to make it stand out as the most preferred destination for the IT companies.

The IT sector did have a significant impact on the Indian economy in the last two decades. There has been a phenomenal growth in the IT and the ITES sectors. Revenues earned through them contribute close to 9.5 percent of India’s GDP of which over 65 percent is earned through foreign exchange. The sector also provides employment to more than 35 lakh professionals directly and one crore people indirectly, making it one of the largest and most attractive private sector employers.
The IT-BPM sector has demonstrated flexibility and as per the Economic Survey 2015-16, it is expected to touch an estimated share of 9.5% of GDP and more than 45% in total services exports in 2015-16. The benefits of IT revolution have started percolating to the hinterland. The Minister for IT, Government of Telangana, K T Rama Rao, said during the launch of the new ICT policy that close to 50 percent of jobs created in this sector have gone to young men and women from the smaller towns and mofussil areas.
Hyderabad has several  congenial factors to further fuel growth in IT sector. It has  earned global reputation as a cosmopolitan city  known for its assimilative cultural ethos. When the IT sector was opening up in the country in mid-90s, Hyderabad became a natural choice for the IT majors. It has become a critical national and international IT hub. The presence of  leading national research and academic instituitions has provided the right eco system for the growth of this knowledge-intensive sector.
Despite challenges in employability of graduates churned out from certain colleges, which resemble as what the Chief Minister himself described as poultry farms, the city  offers the industry with a vast pool of skilled man power vital for the growth of any sector. Hyderabad  has global connectivity. The presence of the world’s major IT companies like Microsoft, Google, Facebook, IBM etc., provide the city a firm launch pad for further momentum in this highly competitive sector.
A large presence of  Telugu disapora who have an emotional connect with the sprawling city  provide the state with avenues for transmission of knowledge through expatriate nationals. According to the Socio Economic Outlook 2016, Government of Telangana, “ICT industry in Telangana State consists of Business Processing Organisations and Knowledge Processing Organisations, providing professional services across the globe. State’s ICT service is mainly oriented towards foreign markets and contributes largely to India’s IT exports.”
Over a period of time, Hyderabad has become a premier global destination for IT and ITES industry of India, next only to Bengaluru. The IT industry in the State has grown by leaps and bounds in recent years. There are 1,300 IT units employing as many as 3.7 lakh professionals in the State, with an annual export turnover of Rs 68,258 crore in 2014-15.
However, in the wake of global economic slowdown, resulting in weak global demand for software exports, companies have adopted various methods such as diversification of market base towards emerging and developing countries, product diversification and cost reduction etc.,  explains  the Socio Economic Outlook, 2016.
Hyderabad accounts  for nearly 11 per cent of national   IT exports. The sector registered  an annual growth rate of 16 per cent in 2014-15 . This is 3 per cent higher than national average growth. The new ICT policy has set ambitious targets. The policy statement aims at making Hyderabad the national leader in terms of IT exports and to leverage on the strong infrastructure that exists to attract more investments.
The objectives of the policy include:  To double the value of current Rs 68,232 crore IT exports in Telangana in five years; to double the current employment of four lakhs in five years; and to establish three tier-2 cities as IT hubs in five years. While Hyderabad will remain the centre of product development and R&D activities, companies, particularly ITES companies, will also be encouraged to locate themselves in tier-2 cities like Warangal, Karimnagar, Nizamabad etc., states the new ICT policy.
Hyderabad already has the premier electronics companies in the public sector. The Electronics Corporation of India Limited (ECIL), Bharat Electronics Limited (BEL) besides defence electronics units provided the much needed industrial base in the city even before the emergence of IT infrastructure. In fact, a large number of semi-skilled persons can get employment in this sector rather than in IT and ITES.
The sub policy on electronics plans to make Hyderabad a global hub for electronics. Two electronics manufacturing clusters will be set up in the suburbs of Hyderabad. These will be dedicated clusters for electronics manufacturing. India home to a new breed of young startups has clearly evolved to become the third largest base of technology startups in the world.  Within the last one year, the number of startups grew by 40%, creating around 80,000 jobs in 2015. This emerging sector is set to get a fillip with the Startup India programme, says the Economic Survey, Government of India, 2015-16.
The new ICT policy states that the Government of Telangana aims to build a booming startup eco system that will unleash the entrepreneurial spirits amongst the youth and in turn aid in job and wealth creation. It aims to bring the entire startup community in the state under one umbrella, T-Hub, located in Hyderabad.
The global gaming and animation market is expected to grow to $240 billion in the next three years.  A significant portion of this work is outsourced to India. Hyderabad is already home to a number of gaming and animation companies. Many of these companies have made a mark in national and global market. The new policy aims to give a further push to it. The gaming and animation sector will further enhance the global brand image for Hyderabad, besides yielding revenue.
Firm foundation, futuristic policy, strategic advantages associated with the city etc., would significantly contribute to the enhancement and enrichment of the IT landscape of Hyderabad. Government, industry, academia etc. should be partners in this Brand Hyderabad mega venture.
HYDERABAD:  After over three years of uncertainty, the Information Technology industry in Hyderabad heaved a sigh of relief with the announcement by the Congress to carve out a separate Telangana state out of Andhra Pradesh.
The announcement has cleared the air over the future of Hyderabad, a major IT destination in the country.
This fast growing metropolis will be the joint capital of Telangana and Andhra Pradesh for 10 years. Andhra Pradesh is expected to build its own capital within that period.
The IT industry welcomed the decision, saying it brought an end to uncertainty and brightened the prospects of the city attracting new investments.
The industry with software exports to the tune of Rs.40,000 crore was worried over frequent shutdowns and spells of violent protests for separate Telangana state. This had taken sheen off brand Hyderabad.
"I believe the decision has brought in a positive sentiment as the uncertainty surrounding the matter has finally been resolved," said V. Laxmikanth, managing director, Broadridge India.
"Hyderabad, known for its large pool of talented IT professionals and as a base for leading MNCs, is expected to reclaim its reputation of being the destination of choice for businesses across industries," he said.
BVR Mohan Reddy, chairman and managing director, Infotech Enterprises, said while he was not for or against the decision, he was anxiously looking forward for this decisive movement.
"The separation problem has been voiced for the last 50 years but in the last three years some of these agitations have been (hurdles) for progress of the state. The uncertainty has slowed down the growth. That is now past."
Suman Reddy, vice president and managing director, Pegasystems India, hoped the development will bring back stability in the business environment of the city.
"Growth and expansion from the perspective of IT MNCs investing in the state had been stalled/relatively slow due to this issue, which hopefully will resume after the decision is announced," he said.
Hyderabad, which emerged on the world IT map only in the mid-1990s, is home to over 1,000 IT and ITeS companies including global majors like Microsoft, Google, Facebook, Dell, Oracle and Amazon.
The IT and ITes sector provides nearly five lakh jobs.
Bengaluru may have hogged the startup spotlight in recent years, but Hyderabad has been working towards building one of the strongest foundations for a powerful startup ecosystem and hub in India. In fact, there was a time when it was believed that Hyderabad would give Bengaluru a tough competition.
At the peak of its IT revolution, Hyderabad was home to biggies like Microsoft, Facebook, and Google. A common anecdote of the time was how then Chief Minister, Chandrababu Naidu, managed a meeting Bill Gates and convinced him to start Microsoft’s Research Centre out of Hyderabad. However, political issues like the Telangana movement put a halt to Hyderabad’s march towards becoming the predominant startup hub.

A group of government officials, investors, incubators and startups in Hyderabad are now attempting to make up for lost time. The results are already visible.  Global e-commerce major Amazon chose the city as the home for its largest warehouse, which is capable of storing about two million products. Uber too is setting up a state-of-the art facility in Hyderabad, which will eventually be its largest international office.
Young and upcoming startups are also getting a boost.  YourStory currently tracks 1,800 startups in Hyderabad.
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Image created by Aditya Ranade
Funding galore
Before we get into the different ingredients, that make a truly powerful Hyderabadi startup biryani, here is a sneak peek into the funding details in the city:
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Funding trend in the past five years in Hyderabad:
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Image created by Aditya Ranade
The Hyderabad Startups
Apart from mobile-app-focussed startups like CanvasFlip, AppVirality and Zify, the health and wellness sector appears to be dominating the scene in terms of funding with the likes of ManageMySpa, Zapluk, MapMyGenome, Truweight and Healthians getting investments. Others include Pricejugaad and MySmartPrice.
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Image Created by Aditya Ranade
In September last year, a NASSCOM report showed that close to eight percent of the startup activity in India is happening in Hyderabad. Sanjay Enishetty, 50K ventures, says that by 2018, the Hyderabad Startup Ecosystem will possibly be bigger than the Bengaluru Startup Ecosystem.
Speaking of the quality and kind of startups that are seen in Hyderabad, Sateesh says that like most startups across India, those in Hyderabad are also jumping on the hyperlocal and e-commerce bandwagon.
However, over the past few years, the kind of startups that have grown in Hyderabad are in the enterprise software, commerce, and healthcare sectors.
The entrepreneurs themselves
Along with the growing number of entrepreneurs and people starting up, the quality of people starting up is also improving. Ramesh says that the quality of entrepreneurship in the city has seen a significant improvement in the past year.
“In Hyderabad, there is this strong trend of people with solid corporate experience quitting their jobs and starting up. This in itself helps improve the quality of entrepreneurship. The entrepreneurs have an understanding of the different elements and aspects of a business. They aren’t necessarily knowledgeable on everything, but they understand that there are different aspects,” says Tom Thomas, COO, CIE Hyderabad.
However, those who have seen Hyderabad grow as a startup hub say that quality of entrepreneurs has always been high. “You look at the top names in the healthcare and pharmaceutical world. The entrepreneurs are from Hyderabad. People from Hyderabad have always been great at entrepreneurship and setting up businesses,” says Sateesh Andra, Managing Director, Endiya Partners.
So what are the key ingredients that make Hyderabad a great startup ecosystem?
I. The infrastructure
For any startup to thrive, apart from the product, you need the right team and infrastructure in place. And Hyderabad has ensured that the city has a strong infrastructural base. In fact, the infrastructural foundation of the city was laid in the early 2000s.
“Hyderabad today can be considered amongst the cheapest metros in the country. The seeds of a strong infrastructural base were sown early on, and now we are just adding to it,” says Ramesh Loganathan, VP Products and Centre head, Progress Software, and one of the city’s most sought-after mentors and advisors.
Building an incubation ecosystem – T-Hub
There possibly has not been a better time for startups in India, with the Startup India initiative and more recent announcement. But with Hyderabad specifically, there has been a surge in government support. Recently, the youngest state in the country, Telangana made headlines with the announcement of its largest incubation centre in India – T-Hub.
It is believed to be one of the most powerful and ambitious projects that has been taken up by the government. K T Rama Rao, who was an IT professional for six years and is now the IT Minister of Telangana, has said that the T-Hub is not for Telangana techies alone, but is also meant to attract talent from across the country.
The Telangana government is also in the process of setting up two more incubation centres in the gaming and animation and aerospace sectors soon Located in the IIIT campus, T-Hub is called ‘the incubator of incubators’ by mentors and incubator heads in Hyderabad.
Explaining this, Ajit Raj of T-Hub says that the aim of the incubation centre is to create a complete ecosystem for startups. “The main idea here is to build a support system that is so strong that startups don’t find it difficult to get advice, mentorship, funding and infrastructure. For that, we not only have our own incubation centre, but also have tie ups with the likes of NASSCOM, CIE and others to help incubate their startups as well,” adds Ajit.
Ratan Tata, when he inaugurated the T-Hub, said, “Walking around the T-Hub building made me realise that we are entering into a new phase of entrepreneurship, innovation and enterprise in India. It is enterprise that is non-traditional and focussed on innovation of the mind. This was what elevated the US in the 80s into a new world of technology and hi-tech. It changed the way we lived. A facility like T-Hub gives you a sneak-peek into the new India of tomorrow.”
Aditya Vuchi, Founder, Zippr, a Hyderabad-based tech startup, says it has never been easier to start up in Telangana. He is hoping for a collaboration between Central and State departments.
The T-Hub, on Monday, also announced its incubation programme LAB/32. In this programme a few handpicked startups will not only be able to access top mentors in the industry but also will be given the opportunity to pitch to VCs and angel investors across the globe. Some of the mentors include Gautham Seshadri of Aarin Capital and Keerti Melkote of Aruba Networks.

The academic incubation centres
“When I was working at the CIE there were close to 11 startups in the centre, and close to a year later, when I was leaving CIE, there were close of 70 startups in the incubation centre,” says Manoj Surya, Founder of Zenty. Apart from CIE in IIIT, the other incubation centres are at ISB and BITS Pilani.
Praveen Dorna, Co-Founder Startup Byte and the Head Incubation manager at CIE says that the kind of entrepreneurs at CIE aren’t very different from entrepreneurs elsewhere, but the idea is that the incubation centre provides a solid base for founders who are looking to build great products and need support of infrastructure, mentors and advisors.
Co-working spaces
Raghuveer Kovuru, Startup Coach and Community Animator at Co.Lab.Orate says that while startup activity in Hyderabad cannot be compared to Bengaluru, the city is nevertheless working slowly and steadily to become a strong ecosystem base.
“The idea of a co-working space is to build community and help startups spend less on office spaces. They focus on building the product and talent base,” says Raghuveer. He adds that there is more collaboration and networking in a co-working space.
In most co-working spaces, there are events and talks on funding, growth, and key advice that any first-time entrepreneur would seek.
II. People
Startups and startup ecosystems are defined by the kind of people who form the startups or the ecosystem. In Hyderabad, with government support and infrastructural elements in place, it is now up to the entrepreneurs to build and develop this ecosystem.
Availability of talent
The growing number of top education institutes in the city not only ensures great incubation centres, but also that startups get the much-needed tech talent and people. Ramesh says that after Bengaluru, Hyderabad possibly has the most thriving talent pool across different sectors. The city has seen growth and development across different verticals, ensuring that the talent has grown to an exponential level in the city.
An interlinked network
Looking at the growing number of entrepreneurs and startups in Hyderabad, key advisors, mentors, incubator heads and investors decided to get together and help entrepreneurs in the city. The idea was simple; it was focussed on helping the startups leverage the experience and understanding of mentors and investors.
Today, Hyderabad is among the most accessible and connected ecosystems in the country. “Everyone is open to talk to and the kind of advice one gets is very open and constructive. This is because, everyone wants the ecosystem to grow and thrive,” adds Manoj.
Sanjay says that when 50k Ventures first started in 2007-08, there weren’t many people to look up to or turn towards for advice. But today, he says that the ecosystem is working as one strong unit helping everyone grow and thrive in their own way.
Stitching the loose ends
While there are different ingredients in place, the Hyderabad startup ecosystem still needs to see the kind of funding that Bengaluru sees. While interconnected and with decent infrastructure in place, the city lost a good part of a year in political conflicts.
Even if there is decent technical skill and talent in the city, startups in Hyderabad will need to compete with Bengaluru-based startups that have deeper pockets. “We have different ingredients in place, we now need to start attracting people from different cities,” adds Tom.
While it might seem that every metro in the country is aiming to be a startup hub, Hyderabad is truly working on different aspects to achieve that end goal. “Everyone talks about the infrastructure, talent and growth opportunity, but it is about using the different elements and truly building a powerful startup ecosystem, and Hyderabad is working tirelessly as one unit towards that end,” says Ramesh.Back in the mid-1990s in the United States, when the Internet boom was just starting, the East was also poised to get in on the technological wagon. Hyderabad, an ancient city borne of mixed traditions and cultures, started the slow transformation into a cyber-city that, today, has become one of India’s premiere information technology and IT-enabled service hubs. Hyderabad has also become the country’s center for scientific and technological development not only in IT infrastructure but also in pharmaceutical and science research industries. To say that Hyderabad is the financial and technological capital of the state of Telangana is not a rumour; it is a fact supported annually by the many international technological firms that call the city their home.
After Bangalore, Hyderabad has been dubbed as the Silicon Valley of India, mainly in light of the fact that more and more firms have set up their operations there. Starting in the 1990s, software companies, business process outsourcing (BPO) firms, call centers, and other technological services were set up, making it the center for all call center operations, technological development, and KPOs in all of India. There are several reasons why these large multinational information technology giants have chosen Hyderabad as one of their base of operations in South Asia: for one, the untapped intellectual talent in the region is immense, especially in the field of technology and mathematics. Another reason is that labour is especially cheap and abundant. Most companies will be able to hire more workers at almost half the cost when they’re in the West. India (and, in extension, Hyderabad) also have labour standards that are pretty much more reasonable and relaxed, as well as a working environment that is conducive to work.
The state’s support for IT sectors is very strong. In Hyderabad, the government’s industrial policy for the IT, Biotechnology, and Pharmaceutical sectors is very liberal and high priority is given to its development especially since it plays a vital role in sustainable economic growth. Extensive investments in the city’s digital infrastructure are also being made to promote construction and setting up of several campuses to house arrays of companies investing in the city’s IT growth. Foremost of these is the development of HITEC City which stands for Hyderabad Information Technology Engineering Consultancy City. It is a major technology township that has become the center of information technology industry in Hyderabad. Composed of several campuses and phases, HITEC City has residential areas where the IT professionals can live, as well as convention centers and malls. Several multinational IT giants have made the City their base of operations in India, including Accenture, Oracle, Verizon, and IBM.
Since Hyderabad offers a stable political climate as well as an efficient and good IT infrastructure, it is safe to assume that it will become a core player in Asia in terms of being information technology hub. The conditions have made Hyderabad a prime destination for offshoring services, making it an alluring destination for many IT companies worldwide.
Telangana Information Technology and Panchayat Raj Minister K.T. Rama Rao has said that the aim of his Government was to drag the preeminent position of Bengaluru in software industry and bring it to Hyderabad.
Noting that the `first mover' advantage in the industry was lost to Bengaluru due to failures of previous governments, Mr. Rao said that the present regime here was focussed on making Hyderabad number one destination of the country in IT sector.
He said at a meet-the-press programme organised by Press Club, Hyderabad, that a huge expansion of IT sector was proposed in Hyderabad whose exports in 2014 were of the order of Rs.57,000 crore, next only to Bengaluru. The goal was to increase exports to Rs.1.20 lakh in collaboration with major companies abroad. The Government would also tap the potential of low-cost and high-value human resource available in Hyderabad by encouraging them to set up startups.
The Telangana Government will start a technology incubator within a fortnight when Chief Minister K. Chandrasekhar Rao will spell out the incentives.
The Minister maintained that the biggest takeaway from his recent US tour was that he succeeded in building confidence of IT industry in Hyderabad as an emerging market. The Government established direct contact with honchos in Silicon Valley. They agreed to participate in IT Hub to be set up in Hyderabad. A separate zone for micro, small and medium enterprises will also be set up over seven acres at Gachibowli.
Information Technology Investment Region (ITIR)
The Information Technology Investment Region (ITIR), Hyderabad is an upcoming IT investment region jointly being developed by Government of India and Government of Telangana.[44]  The Union government has given in-principle approval on Sept 8th, 2012 for the Information Technology Investment Region (ITIR) for development of self-contained integrated knowledge clusters for growth of IT and electronic hardware manufacturing in 50,000 acres in and around Hyderabad.[45]  The project which is modeled along the lines of Shenzhen SEZ in China, is aimed at attracting an investment of Rs 2.19 trillion ($44 billion) in the IT, ITES and electronics sectors and providing direct employment to 1.5 million people.
Under the mega project, special economic zones (SEZs), industrial parks, free trade zones, warehousing zones and export-oriented units would come up in three corridors around the city which includes the areas Madhapur, Gachibowli, Uppal, Mamidipalli, Raviryal, Adibatla and Maheswaram, and Pocharam. The Government proposed to develop the infrastructure for ITIR at an estimated cost of over Rs 2.19 lakh crore.
On September 20, 2013 the Central Government gave the official approval for the ITIR project[46]
In April 2016, the Central Government informed Telangana state Government that it would revise the entire plan. This, according to sources, may lead to the delay in the estimated investments of Rs 3 lakh crore and creation of nearly 60 lakh jobs. No explanation given on why the need for revise plan.

Which state in the U.S. has the largest information technology industry?