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96-500: Durum wheat ( / ˈ dj ʊər ə m / ), also called pasta wheat or macaroni wheat ( Triticum durum or Triticum turgidum subsp. durum ), is a tetraploid species of wheat . It is the second most cultivated species of wheat after common wheat , although it represents only 5% to 8% of global wheat production. It was developed by artificial selection of the domesticated emmer wheat strains formerly grown in Central Europe and

192-409: A Pneumocystis carinii infection, which indicates a weak immune system. Triploidy may be the result of either digyny (the extra haploid set is from the mother) or diandry (the extra haploid set is from the father). Diandry is mostly caused by reduplication of the paternal haploid set from a single sperm, but may also be the consequence of dispermic (two sperm) fertilization of the egg. Digyny

288-626: A hybrid genome with two sets of chromosomes derived from Triticum urartu and two sets of chromosomes derived from Aegilops speltoides . Each chromosome pair derived from the Triticum urartu parent is homoeologous to the opposite chromosome pair derived from the Aegilops speltoides parent, though each chromosome pair unto itself is homologous . Examples in animals are more common in non-vertebrates such as flatworms , leeches , and brine shrimp . Within vertebrates, examples of stable polyploidy include

384-410: A miscarriage ; those that do survive to term typically die shortly after birth. In some cases, survival past birth may be extended if there is mixoploidy with both a diploid and a triploid cell population present. There has been one report of a child surviving to the age of seven months with complete triploidy syndrome. He failed to exhibit normal mental or physical neonatal development, and died from

480-476: A few green olives", not to be confused with Neapolitan pizza ; calabresa , with slices of longaniza ; jamon y morrones , with sliced ham and roasted bell peppers ; as well as versions with provolone , with anchovies , with hearts of palm , or with chopped hard boiled egg . A typical custom that is unique to Buenos Aires is to accompany pizza with fainá , a pancake made from chickpea flour . The terms dessert pizza and sweet pizza are used for

576-596: A high temperature, traditionally in a wood-fired oven . The term pizza was first recorded in the year 997 AD, in a Latin manuscript from the southern Italian town of Gaeta , in Lazio, on the border with Campania . Raffaele Esposito is often credited for creating the modern pizza in Naples . In 2009, Neapolitan pizza was registered with the European Union as a traditional speciality guaranteed (TSG) dish. In 2017,

672-414: A level of diversity intermediate between those groups. There is evidence of an increase in the intensity of breeding after 1990. Commercially produced dry pasta , or pasta secca , is made almost exclusively from durum semolina . Most home-made fresh pastas also use durum wheat or a combination of soft and hard wheats. Husked but unground, or coarsely ground, it is used to produce the semolina in

768-477: A meal that includes round cakes (such as pita bread) topped with cooked vegetables. When they eat the bread, they realize that these are the "tables" prophesied by Celaeno. In 2023, archeologists discovered a fresco in Pompeii appearing to depict a pizza-like dish among other foodstuffs and staples on a silver platter. Italy's culture minister said it "may be a distant ancestor of the modern dish". The first mention of

864-846: A mechanism for a kind of 'reverse speciation', whereby gene flow is enabled following the polyploidy event, even between lineages that previously experienced no gene flow as diploids. This has been detailed at the genomic level in Arabidopsis arenosa and Arabidopsis lyrata . Each of these species experienced independent autopolyploidy events (within-species polyploidy, described below), which then enabled subsequent interspecies gene flow of adaptive alleles, in this case stabilising each young polyploid lineage. Such polyploidy-enabled adaptive introgression may allow polyploids at act as 'allelic sponges', whereby they accumulate cryptic genomic variation that may be recruited upon encountering later environmental challenges. Polyploid types are labeled according to

960-481: A more common path to allopolyploidy because F 1 hybrids between taxa often have relatively high rates of unreduced gamete formation – divergence between the genomes of the two taxa result in abnormal pairing between homoeologous chromosomes or nondisjunction during meiosis. In this case, allopolyploidy can actually restore normal, bivalent meiotic pairing by providing each homoeologous chromosome with its own homologue. If divergence between homoeologous chromosomes

1056-403: A paleopolyploid. It is a middle aged polyploid. Often this refers to whole genome duplication followed by intermediate levels of diploidization. Ancient genome duplications probably occurred in the evolutionary history of all life. Duplication events that occurred long ago in the history of various evolutionary lineages can be difficult to detect because of subsequent diploidization (such that

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1152-436: A pizza can be baked on a pizza stone in a regular oven to reproduce some of the heating effect of a brick oven. Cooking directly on a metal surface results in too rapid heat transfer to the crust, burning it. Some home chefs use a wood-fired pizza oven, usually installed outdoors. As in restaurants, these are often dome-shaped, as pizza ovens have been for centuries, in order to achieve even heat distribution. Another variation

1248-695: A polyploid starts to behave cytogenetically as a diploid over time) as mutations and gene translations gradually make one copy of each chromosome unlike the other copy. Over time, it is also common for duplicated copies of genes to accumulate mutations and become inactive pseudogenes. In many cases, these events can be inferred only through comparing sequenced genomes . Examples of unexpected but recently confirmed ancient genome duplications include baker's yeast ( Saccharomyces cerevisiae ), mustard weed/thale cress ( Arabidopsis thaliana ), rice ( Oryza sativa ), and two rounds of whole genome duplication (the 2R hypothesis ) in an early evolutionary ancestor of

1344-614: A small-sized, thick-crust, deep-dish pizza typically served in Turin , Piedmont. The first pizzeria in the US was opened in New York City's Little Italy in 1905. Common toppings for pizza in the United States include anchovies , ground beef, chicken, ham, mushrooms, olives, onions, peppers, pepperoni , pineapple, salami, sausage, spinach, steak, and tomatoes. Distinct regional types developed in

1440-579: A sterile triploid hybrid between E. guttata and E. lutea, both of which have been introduced and naturalised in the United Kingdom. New populations of E. peregrina arose on the Scottish mainland and the Orkney Islands via genome duplication from local populations of E. × robertsii . Because of a rare genetic mutation, E. peregrina is not sterile. On the other hand, polyploidization can also be

1536-413: A total of 28, unlike hard red winter and hard red spring wheats , which are hexaploid (six sets of chromosomes) for a total of 42. Durum wheat originated through intergeneric hybridization and polyploidization involving two diploid (having two sets of chromosomes) grass species: T. urartu (2n=2x=14, AA genome) and a B-genome diploid related to Aegilops speltoides (2n=2x=14, SS genome) and

1632-499: A variety of dishes resembling a pizza, including chocolate pizza and fruit pizza . Some are based on a traditional yeast dough pizza base, while others have a cookie -like base and resemble a traditional pizza solely in having a flat round shape with a distinct base and topping. Some pizza restaurants offer dessert pizzas. Some pizzas mass-produced by American pizza chains have been criticized as having an unhealthy balance of ingredients. Pizza can be high in salt and fat, and

1728-647: Is Sicilian pizza (locally called sfincione or sfinciuni ), a thick-crust or deep-dish pizza originating during the 17th century in Sicily : it is essentially a focaccia that is typically topped with tomato sauce and other ingredients. Until the 1860s, sfincione was the type of pizza usually consumed in Sicily, especially in the Western portion of the island. Other variations of pizzas are also found in other regions of Italy, for example pizza al padellino or pizza al tegamino ,

1824-567: Is a non-profit organization founded in 1984 with headquarters in Naples that aims to promote traditional Neapolitan pizza. In 2009, upon Italy's request, Neapolitan pizza was registered with the European Union as a traditional speciality guaranteed (TSG) dish, and in 2017 the art of its making was included on UNESCO 's list of intangible cultural heritage . Pizza is sold fresh or frozen , and whole or in portion-size slices . Methods have been developed to overcome challenges such as preventing

1920-411: Is a mainstay of the country's cuisine , especially of its capital Buenos Aires , where it is regarded as a cultural heritage and icon of the city. Argentina is the country with the most pizzerias per inhabitant in the world and, although they are consumed throughout the country, the highest concentration of pizzerias and customers is Buenos Aires, the city with the highest consumption of pizzas in

2016-452: Is also one of the only flours that is purposely oxidized for flavor and color. There is also a red durum, used mostly for livestock feed . The cultivation of durum generates greater yield than other wheats in areas of low precipitation. Good yields can be obtained by irrigation , but this is rarely done. In the first half of the 20th century, the crop was widely grown in Russia . Durum is one of

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2112-407: Is common among ferns and flowering plants (see Hibiscus rosa-sinensis ), including both wild and cultivated species . Wheat , for example, after millennia of hybridization and modification by humans, has strains that are diploid (two sets of chromosomes), tetraploid (four sets of chromosomes) with the common name of durum or macaroni wheat, and hexaploid (six sets of chromosomes) with

2208-446: Is dotted by past and recent whole-genome duplication events (see Albertin and Marullo 2012 for review). Several examples of polyploids are known: In addition, polyploidy is frequently associated with hybridization and reticulate evolution that appear to be highly prevalent in several fungal taxa. Indeed, homoploid speciation (hybrid speciation without a change in chromosome number) has been evidenced for some fungal species (such as

2304-449: Is eaten with the use of a knife and fork. In casual settings, however, it is typically cut into slices to be eaten while held in the hand . Pizza is also sold in grocery stores in a variety of forms, including frozen or as kits for self-assembly. Store-bought pizzas are then cooked using a home oven . In 2017, the world pizza market was US$ 128 billion, and in the US it was $ 44 billion spread over 76,000 pizzerias. Overall, 13% of

2400-698: Is even across the two subgenomes, this can theoretically result in rapid restoration of bivalent pairing and disomic inheritance following allopolyploidization. However multivalent pairing is common in many recently formed allopolyploids, so it is likely that the majority of meiotic stabilization occurs gradually through selection. Because pairing between homoeologous chromosomes is rare in established allopolyploids, they may benefit from fixed heterozygosity of homoeologous alleles. In certain cases, such heterozygosity can have beneficial heterotic effects, either in terms of fitness in natural contexts or desirable traits in agricultural contexts. This could partially explain

2496-488: Is even imported. On the other hand, many countries in Europe produce durum in commercially significant quantities. In India durum accounts for roughly 5% of total wheat production in the country, and is used to make products such as rava and sooji. Durum wheat is subject to four processes: cleaning, tempering, milling and purifying. First, durum wheat is cleaned to remove foreign material and shrunken and broken kernels. Then it

2592-885: Is frequent in plants, some estimates suggesting that 30–80% of living plant species are polyploid, and many lineages show evidence of ancient polyploidy ( paleopolyploidy ) in their genomes. Huge explosions in angiosperm species diversity appear to have coincided with the timing of ancient genome duplications shared by many species. It has been established that 15% of angiosperm and 31% of fern speciation events are accompanied by ploidy increase. Polyploid plants can arise spontaneously in nature by several mechanisms, including meiotic or mitotic failures, and fusion of unreduced (2 n ) gametes. Both autopolyploids (e.g. potato ) and allopolyploids (such as canola, wheat and cotton) can be found among both wild and domesticated plant species. Most polyploids display novel variation or morphologies relative to their parental species, that may contribute to

2688-422: Is grilled pizza, in which the pizza is baked directly on a barbecue grill. Some types, such as Sicilian pizza , deep-dish Chicago-style pizza and Greek pizza , are baked in a pan rather than directly on the bricks of the pizza oven. Most restaurants use standard and purpose-built pizza preparation tables to assemble their pizzas. Mass production of pizza by chains can be completely automated. The bottom of

2784-416: Is highly resistant to such exposures. The mechanism by which the genome is accurately restored involves RecA-mediated homologous recombination and a process referred to as extended synthesis-dependent strand annealing (SDSA) . Pizza Pizza is an Italian dish typically consisting of a flat base of leavened wheat-based dough topped with tomato , cheese , and other ingredients, baked at

2880-514: Is more rarely diagnosed than triploidy, but is observed in 1–2% of early miscarriages. However, some tetraploid cells are commonly found in chromosome analysis at prenatal diagnosis and these are generally considered 'harmless'. It is not clear whether these tetraploid cells simply tend to arise during in vitro cell culture or whether they are also present in placental cells in vivo . There are, at any rate, very few clinical reports of fetuses/infants diagnosed with tetraploidy mosaicism. Mixoploidy

2976-473: Is most commonly caused by either failure of one meiotic division during oogenesis leading to a diploid oocyte or failure to extrude one polar body from the oocyte . Diandry appears to predominate among early miscarriages , while digyny predominates among triploid zygotes that survive into the fetal period. However, among early miscarriages, digyny is also more common in those cases less than 8 + 1 ⁄ 2 weeks gestational age or those in which an embryo

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3072-433: Is most commonly induced by treating seeds with the chemical colchicine . Some crops are found in a variety of ploidies: tulips and lilies are commonly found as both diploid and triploid; daylilies ( Hemerocallis cultivars) are available as either diploid or tetraploid; apples and kinnow mandarins can be diploid, triploid, or tetraploid. Besides plants and animals, the evolutionary history of various fungal species

3168-407: Is not a strong wheat in the sense of giving strength to dough through the formation of a gluten network. Durum contains 27% extractable wet gluten, about 3% higher than common wheat ( T. aestivum L.). Some authorities synonymize "durum" and Triticum turgidum . Some reserve "durum" for Triticum turgidum subsp. durum . Durum wheat is a tetraploid wheat, having four sets of chromosomes for

3264-623: Is often associated with apomictic mating systems. In agricultural systems, autotriploidy can result in seedlessness, as in watermelons and bananas . Triploidy is also utilized in salmon and trout farming to induce sterility. Rarely, autopolyploids arise from spontaneous, somatic genome doubling, which has been observed in apple ( Malus domesticus ) bud sports . This is also the most common pathway of artificially induced polyploidy, where methods such as protoplast fusion or treatment with colchicine , oryzalin or mitotic inhibitors are used to disrupt normal mitotic division, which results in

3360-440: Is part of cytology and, more specifically, cytogenetics . Although the replication and transcription of DNA is highly standardized in eukaryotes , the same cannot be said for their karyotypes, which are highly variable between species in chromosome number and in detailed organization despite being constructed out of the same macromolecules. In some cases, there is even significant variation within species. This variation provides

3456-435: Is present. There are also two distinct phenotypes in triploid placentas and fetuses that are dependent on the origin of the extra haploid set. In digyny, there is typically an asymmetric poorly grown fetus , with marked adrenal hypoplasia and a very small placenta . In diandry, a partial hydatidiform mole develops. These parent-of-origin effects reflect the effects of genomic imprinting . Complete tetraploidy

3552-412: Is quite commonly observed in human preimplantation embryos and includes haploid/diploid as well as diploid/tetraploid mixed cell populations. It is unknown whether these embryos fail to implant and are therefore rarely detected in ongoing pregnancies or if there is simply a selective process favoring the diploid cells. A polyploidy event occurred within the stem lineage of the teleost fish. Polyploidy

3648-471: Is sometimes called pizzetta . A person who makes pizza is known as a pizzaiolo . The word pizza was borrowed from Italian into English in the 1930s; before it became well known, pizza was called "tomato pie" by English speakers. Some regional pizza variations still use the name tomato pie. Records of pizza-like foods can be found throughout ancient history. In the 6th century BC, the Persian soldiers of

3744-423: Is tempered to a moisture content, toughening the seed coat for efficient separation of bran and endosperm. Durum milling is a complex procedure involving repetitive grinding and sieving . Proper purifying results in maximum semolina yield and the least amount of bran powder . To produce bread, durum wheat is ground into flour. The flour is mixed with water to produce dough. The quantities mixed vary, depending on

3840-624: Is the Coast Redwood Sequoia sempervirens , which is a hexaploid (6 x ) with 66 chromosomes (2 n = 6 x = 66), although the origin is unclear. Aquatic plants, especially the Monocotyledons , include a large number of polyploids. The induction of polyploidy is a common technique to overcome the sterility of a hybrid species during plant breeding. For example, triticale is the hybrid of wheat ( Triticum turgidum ) and rye ( Secale cereale ). It combines sought-after characteristics of

3936-437: Is thus an allotetraploid (having four sets of chromosomes, from unlike parents) species. Durum—and indeed all tetraploids —lack Fhb1 alleles . The only exception is found by Buerstmayr et al. , 2012 on the 3B chromosome . One of the predominant production areas of durum—Italy—has domesticated varieties with lower genetic diversity than wild types, but ssp. turanicum , ssp. polonicum and ssp. carthlicum have

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4032-483: Is uncertain when tomatoes were first added and there are many conflicting claims, although it certainly could not have been before the 16th century and the Columbian Exchange . Pizza was sold from open-air stands and out of pizza bakeries until about 1830, when pizzerias in Naples started to have stanze with tables where clients could sit and eat their pizzas on the spot. A popular contemporary legend holds that

4128-551: Is under- or over-represented are said to be aneuploid (from the Greek words meaning "not", "good", and "fold"). Aneuploidy refers to a numerical change in part of the chromosome set, whereas polyploidy refers to a numerical change in the whole set of chromosomes. Polyploidy occurs in some tissues of animals that are otherwise diploid, such as human muscle tissues. This is known as endopolyploidy . Species whose cells do not have nuclei, that is, prokaryotes , may be polyploid, as seen in

4224-528: Is unsuitable for people with gluten-related disorders such as celiac disease , non-celiac gluten sensitivity and wheat allergy . Tetraploid Polyploidy is the result of whole-genome duplication during the evolution of species. It may occur due to abnormal cell division , either during mitosis, or more commonly from the failure of chromosomes to separate during meiosis or from the fertilization of an egg by more than one sperm. In addition, it can be induced in plants and cell cultures by some chemicals:

4320-737: The Achaemenid Empire during the rule of Darius the Great baked flatbreads with cheese and dates on top of their battle shields and the ancient Greeks supplemented their bread with oils , herbs , and cheese . An early reference to a pizza-like food occurs in the Aeneid , when Celaeno, queen of the Harpies , foretells that the Trojans would not find peace until they are forced by hunger to eat their tables (Book III). In Book VII, Aeneas and his men are served

4416-793: The Near East around 7000 BC, which developed a naked, free-threshing form. Like emmer, durum wheat is awned (with bristles). It is the predominant wheat that grows in the Middle East. Durum in Latin means 'hard', and the species is the hardest of all wheats. This refers to the resistance of the grain to milling, in particular of the starchy endosperm , causing dough made from its flour to be weak or "soft". This makes durum favorable for semolina and pasta and less practical for flour, which requires more work than with hexaploid wheats such as common bread wheats . Despite its high protein content, durum

4512-546: The basidiomycota Microbotryum violaceum ). As for plants and animals, fungal hybrids and polyploids display structural and functional modifications compared to their progenitors and diploid counterparts. In particular, the structural and functional outcomes of polyploid Saccharomyces genomes strikingly reflect the evolutionary fate of plant polyploid ones. Large chromosomal rearrangements leading to chimeric chromosomes have been described, as well as more punctual genetic modifications such as gene loss. The homoealleles of

4608-631: The couscous of North Africa and the Levant . It is also used for Levantine dishes such as tabbouleh , kashk , kibbeh , bitfun and the bulgur for pilafs . In North African cuisine and Levantine cuisine , it forms the basis of many soups , gruels, stuffings, puddings and pastries . When ground as fine as flour, it is used for making bread . In the Middle East , it is used for flat round breads , and in Europe and elsewhere, it can be used for pizza or torte . The use of wheat to produce pasta

4704-664: The salmonids and many cyprinids (i.e. carp ). Some fish have as many as 400 chromosomes. Polyploidy also occurs commonly in amphibians; for example the biomedically important genus Xenopus contains many different species with as many as 12 sets of chromosomes (dodecaploid). Polyploid lizards are also quite common. Most are sterile and reproduce by parthenogenesis ; others, like Liolaemus chiliensis , maintain sexual reproduction. Polyploid mole salamanders (mostly triploids) are all female and reproduce by kleptogenesis , "stealing" spermatophores from diploid males of related species to trigger egg development but not incorporating

4800-420: The vertebrates (which includes the human lineage) and another near the origin of the teleost fishes . Angiosperms ( flowering plants ) have paleopolyploidy in their ancestry. All eukaryotes probably have experienced a polyploidy event at some point in their evolutionary history. A karyotype is the characteristic chromosome complement of a eukaryote species . The preparation and study of karyotypes

4896-538: The 1930s–1940s. Pizza was taken to the United States by Italian immigrants in the late 19th century and first appeared in areas where they concentrated. The country's first pizzeria, Lombardi's , opened in New York City in 1905. Italian Americans migrating from East to West brought with them their dish. From there, the American version exported to the rest of the world. The Associazione Verace Pizza Napoletana ( lit.   ' True Neapolitan Pizza Association ' )

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4992-610: The 20th century, including Buffalo , California , Chicago , Detroit , Greek , New Haven , New York , and St. Louis styles. These regional variations include deep-dish, stuffed, pockets, turnovers, rolled , and pizza-on-a-stick, each with seemingly limitless combinations of sauce and toppings. Thirteen percent of the United States population consumes pizza on any given day. Pizza chains such as Domino's Pizza , Pizza Hut , and Papa John's , pizzas from take and bake pizzerias , and chilled or frozen pizzas from supermarkets make pizza readily available nationwide. Argentine pizza

5088-621: The Argentine capital. The impoverished Italian immigrants that arrived to the city transformed the originally modest dish into a much more hefty meal, motivated by the abundance of food in Argentina. In the 1930s, pizza was cemented as a cultural icon in Buenos Aires, with the new pizzerias becoming a central space for sociability for the working class people who flocked to the city. The most characteristic style of Argentine pizza—which almost all

5184-539: The US population aged two years and over consumed pizza on any given day. The oldest recorded usage of the word pizza is from a Latin text from the town of Gaeta , then still part of the Byzantine Empire , in 997 AD; the text states that a tenant of certain property is to give the bishop of Gaeta duodecim pizze ( lit.   ' twelve pizzas ' ) every Christmas Day , and another twelve every Easter Sunday . Suggested etymologies include: A small pizza

5280-566: The acidity of the mixture. To produce fluffy bread, the dough is mixed with yeast and lukewarm water, heavily kneaded to form a gas-retaining gluten network, and then fermented for hours, producing CO 2 bubbles. The quality of the bread produced depends on the viscoelastic properties of gluten, the protein content and protein composition. Containing about 12% total protein in defatted flour compared to 11% in common wheat, durum wheat yields 27% extractable, wet gluten compared to 24% in common wheat. Because durum wheat contains gluten , it

5376-425: The agamic complexes of Crepis . Some plants are triploid. As meiosis is disturbed, these plants are sterile, with all plants having the same genetic constitution: Among them, the exclusively vegetatively propagated saffron crocus ( Crocus sativus ). Also, the extremely rare Tasmanian shrub Lomatia tasmanica is a triploid sterile species. There are few naturally occurring polyploid conifers . One example

5472-575: The allotetraploid yeast S. pastorianus show unequal contribution to the transcriptome . Phenotypic diversification is also observed following polyploidization and/or hybridization in fungi, producing the fuel for natural selection and subsequent adaptation and speciation. Other eukaryotic taxa have experienced one or more polyploidization events during their evolutionary history (see Albertin and Marullo, 2012 for review). The oomycetes , which are non-true fungi members, contain several examples of paleopolyploid and polyploid species, such as within

5568-613: The archetypal pizza, pizza Margherita , was invented in 1889, when the Royal Palace of Capodimonte commissioned the Neapolitan pizzaiolo ( ' pizza maker ' ) Raffaele Esposito to create a pizza in honor of the visiting Queen Margherita . Of the three different pizzas he created, the queen strongly preferred a pizza swathed in the colors of the Italian flag —red (tomato), green (basil), and white (mozzarella). Supposedly, this type of pizza

5664-404: The art of making Neapolitan pizza was included on UNESCO 's list of intangible cultural heritage . Pizza and its variants are among the most popular foods in the world. Pizza is sold at a variety of restaurants, including pizzerias (pizza specialty restaurants), Mediterranean restaurants , via delivery , and as street food . In Italy, pizza served in a restaurant is presented unsliced, and

5760-567: The basis for a range of studies in what might be called evolutionary cytology. Homoeologous chromosomes are those brought together following inter-species hybridization and allopolyploidization , and whose relationship was completely homologous in an ancestral species. For example, durum wheat is the result of the inter-species hybridization of two diploid grass species Triticum urartu and Aegilops speltoides . Both diploid ancestors had two sets of 7 chromosomes, which were similar in terms of size and genes contained on them. Durum wheat contains

5856-432: The best known is colchicine , which can result in chromosome doubling, though its use may have other less obvious consequences as well. Oryzalin will also double the existing chromosome content. Among mammals , a high frequency of polyploid cells is found in organs such as the brain, liver, heart, and bone marrow. It also occurs in the somatic cells of other animals , such as goldfish , salmon , and salamanders . It

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5952-421: The choice of toppings and sometimes also crust. There are also several styles of pizza, defined by their preparation method. The following lists feature only the notable ones. Authentic Neapolitan pizza (Italian: pizza napoletana ) is made with San Marzano tomatoes , grown on the volcanic plains south of Mount Vesuvius , and either mozzarella di bufala campana , made with milk from water buffalo raised in

6048-474: The classic pizzerias in Buenos Aires specialize in—is the so-called pizza de molde (Spanish for 'pizza in the pan'), characterized by having a "thick, spongy base and elevated bready crust". This style, which today is identified as the typical style of Argentine pizza—characterized by a thick crust and a large amount of cheese—arose when impoverished Italian immigrants found a greater abundance of food in then-prosperous Argentina, which motivated them to transform

6144-457: The common name of bread wheat. Many agriculturally important plants of the genus Brassica are also tetraploids. Sugarcane can have ploidy levels higher than octaploid . Polyploidization can be a mechanism of  sympatric speciation  because polyploids are usually unable to interbreed with their diploid ancestors. An example is the plant Erythranthe peregrina . Sequencing confirmed that this species originated from E. × robertsii ,

6240-745: The crust. Dipping sauce specifically for pizza was invented by American pizza chain Papa John's Pizza in 1984 and has since been adopted by some when eating pizza, especially the crust. Mozzarella is commonly used on pizza, with the buffalo mozzarella produced in the surroundings of Naples. Other cheeses are also used, particularly burrata , Gorgonzola , provolone , pecorino romano , ricotta , and scamorza . Less expensive processed cheeses or cheese analogues have been developed for mass-market pizzas to produce desirable qualities such as browning, melting, stretchiness, consistent fat and moisture content, and stable shelf life . This quest to create

6336-407: The diploid species. A similar relationship exists between three diploid species of Tragopogon ( T. dubius , T. pratensis , and T. porrifolius ) and two allotetraploid species ( T. mirus and T. miscellus ). Complex patterns of allopolyploid evolution have also been observed in animals, as in the frog genus Xenopus . Organisms in which a particular chromosome, or chromosome segment,

6432-402: The durum grown today is amber durum , the grains of which are amber-colored due to the extra carotenoid pigments and are larger than those of other types of wheat. Durum has a yellow endosperm , which gives pasta its color. When durum is milled , the endosperm is ground into a granular product called semolina . Semolina made from durum is used for premium pastas and breads . Notably semolina

6528-581: The ends of a spectrum of divergence between parental subgenomes. Polyploids that fall between these two extremes, which are often referred to as segmental allopolyploids, may display intermediate levels of polysomic inheritance that vary by locus. About half of all polyploids are thought to be the result of autopolyploidy, although many factors make this proportion hard to estimate. Allopolyploids or amphipolyploids or heteropolyploids are polyploids with chromosomes derived from two or more diverged taxa. As in autopolyploidy, this primarily occurs through

6624-471: The evolved polyploids. The high degree of homology among duplicated chromosomes causes autopolyploids to display polysomic inheritance . This trait is often used as a diagnostic criterion to distinguish autopolyploids from allopolyploids, which commonly display disomic inheritance after they progress past the neopolyploid stage. While most polyploid species are unambiguously characterized as either autopolyploid or allopolyploid, these categories represent

6720-604: The four expected if it were truly a tetraploid. This rodent is not a rat , but kin to guinea pigs and chinchillas . Its "new" diploid (2 n ) number is 102 and so its cells are roughly twice normal size. Its closest living relation is Octomys mimax , the Andean Viscacha-Rat of the same family, whose 2 n = 56. It was therefore surmised that an Octomys -like ancestor produced tetraploid (i.e., 2 n = 4 x = 112) offspring that were, by virtue of their doubled chromosomes, reproductively isolated from their parents. Polyploidy

6816-436: The frog, Xenopus (an extension of the work of Briggs and King in 1952) that were able to develop to the tadpole stage. The British scientist J. B. S. Haldane hailed the work for its potential medical applications and, in describing the results, became one of the first to use the word " clone " in reference to animals. Later work by Shinya Yamanaka showed how mature cells can be reprogrammed to become pluripotent, extending

6912-425: The fusion of unreduced (2 n ) gametes, which can take place before or after hybridization . In the former case, unreduced gametes from each diploid taxon – or reduced gametes from two autotetraploid taxa – combine to form allopolyploid offspring. In the latter case, one or more diploid F 1 hybrids produce unreduced gametes that fuse to form allopolyploid progeny. Hybridization followed by genome duplication may be

7008-509: The fusion of unreduced (2 n ) gametes, which results in either triploid ( n + 2 n = 3 n ) or tetraploid (2 n + 2 n = 4 n ) offspring. Triploid offspring are typically sterile (as in the phenomenon of triploid block ), but in some cases they may produce high proportions of unreduced gametes and thus aid the formation of tetraploids. This pathway to tetraploidy is referred to as the triploid bridge . Triploids may also persist through asexual reproduction . In fact, stable autotriploidy in plants

7104-648: The genus Phytophthora . Some species of brown algae ( Fucales , Laminariales and diatoms ) contain apparent polyploid genomes. In the Alveolata group, the remarkable species Paramecium tetraurelia underwent three successive rounds of whole-genome duplication and established itself as a major model for paleopolyploid studies. Each Deinococcus radiodurans bacterium contains 4-8 copies of its chromosome . Exposure of D. radiodurans to X-ray irradiation or desiccation can shatter its genomes into hundred of short random fragments. Nevertheless, D. radiodurans

7200-438: The heat source, an electric deck oven, a conveyor belt oven, or, in traditional style in a wood or coal-fired brick oven . The pizza is slid into the oven on a long paddle, called " peel ", and baked directly on hot bricks, a screen (a round metal grate, typically aluminum), or whatever the oven surface is. Before use, a peel is typically sprinkled with cornmeal to allow the pizza to easily slide on and off it. When made at home,

7296-415: The iconic fugazza and its derivative fugazzeta or fugazza con queso (a terminology that varies depending on the pizzeria), or the pizza de cancha or canchera (a cheese-less variant). Most pizza menus include standard flavor combinations, including the traditional plain mozzarella, nicknamed " muza " or " musa "; the napolitana or " napo ", with "cheese, sliced tomatoes, garlic, dried oregano and

7392-458: The ideal and economical pizza cheese has involved many studies and experiments analyzing the impact of vegetable oil , manufacturing and culture processes, denatured whey proteins, and other changes in manufacture. In 1997, it was estimated that annual production of pizza cheese was 1 million metric tons (1,100,000 short tons) in the US and 100,000 metric tons (110,000 short tons) in Europe. A great number of pizza varieties exist, defined by

7488-429: The large bacterium Epulopiscium fishelsoni . Hence ploidy is defined with respect to a cell. A monoploid has only one set of chromosomes and the term is usually only applied to cells or organisms that are normally diploid. The more general term for such organisms is haploid . A polyploid that is newly formed. That has become polyploid in more recent history; it is not as new as a neopolyploid and not as old as

7584-518: The males' DNA into the offspring. While some tissues of mammals, such as parenchymal liver cells, are polyploid, rare instances of polyploid mammals are known, but most often result in prenatal death. An octodontid rodent of Argentina 's harsh desert regions, known as the plains viscacha rat ( Tympanoctomys barrerae ) has been reported as an exception to this 'rule'. However, careful analysis using chromosome paints shows that there are only two copies of each chromosome in T. barrerae , not

7680-525: The marshlands of Campania and Lazio , or fior di latte . Buffalo mozzarella is protected with its own European protected designation of origin (PDO). Other traditional pizzas include pizza marinara , supposedly the most ancient tomato-topped pizza, pizza capricciosa , which is prepared with mozzarella cheese, baked ham, mushroom, artichoke, and tomato, and pizza pugliese , prepared with tomato, mozzarella, and onions. A popular variant of pizza in Italy

7776-502: The most important food crops in West Asia . Although the variety of the wheat there is diverse, it is not extensively grown there, and thus must be imported. West amber durum produced in Canada is used mostly as semolina/pasta, but some is also exported to Italy for bread production. In the Middle East and North Africa, local bread-making accounts for half the consumption of durum. Some flour

7872-408: The number of chromosome sets in the nucleus . The letter x is used to represent the number of chromosomes in a single set: Autopolyploids are polyploids with multiple chromosome sets derived from a single taxon . Two examples of natural autopolyploids are the piggyback plant, Tolmiea menzisii and the white sturgeon, Acipenser transmontanum . Most instances of autopolyploidy result from

7968-488: The originally modest dish into a much more hefty meal suitable for a main course . The name pizza de molde emerged because there were no pizza ovens in the city, so bakers resorted to baking them in pans. Since they used bakery plates, Argentine pizzas were initially square or rectangular, a format associated with the 1920s that is still maintained in some classic pizzerias, especially for vegetable pizzas, fugazzetas or fugazzas . Other styles of Argentine pizza include

8064-417: The parents, but the initial hybrids are sterile. After polyploidization, the hybrid becomes fertile and can thus be further propagated to become triticale. In some situations, polyploid crops are preferred because they are sterile. For example, many seedless fruit varieties are seedless as a result of polyploidy. Such crops are propagated using asexual techniques, such as grafting . Polyploidy in crop plants

8160-414: The pizza, called the "crust", may vary widely according to style—thin as in a typical hand-tossed Neapolitan pizza or thick as in a deep-dish Chicago-style . It is traditionally plain, but may also be seasoned with garlic or herbs, or stuffed with cheese. The outer edge of the pizza is sometimes referred to as the cornicione . Some pizza dough contains sugar, to help its yeast rise and enhance browning of

8256-715: The possibilities to non-stem cells. Gurdon and Yamanaka were jointly awarded the Nobel Prize in 2012 for this work. True polyploidy rarely occurs in humans, although polyploid cells occur in highly differentiated tissue, such as liver parenchyma , heart muscle, placenta and in bone marrow. Aneuploidy is more common. Polyploidy occurs in humans in the form of triploidy , with 69 chromosomes (sometimes called 69, XXX), and tetraploidy with 92 chromosomes (sometimes called 92, XXXX). Triploidy, usually due to polyspermy , occurs in about 2–3% of all human pregnancies and ~15% of miscarriages. The vast majority of triploid conceptions end as

8352-545: The prevalence of allopolyploidy among crop species. Both bread wheat and triticale are examples of an allopolyploids with six chromosome sets. Cotton , peanut , and quinoa are allotetraploids with multiple origins. In Brassicaceous crops, the Triangle of U describes the relationships between the three common diploid Brassicas ( B. oleracea , B. rapa , and B. nigra ) and three allotetraploids ( B. napus , B. juncea , and B. carinata ) derived from hybridization among

8448-796: The processes of speciation and eco-niche exploitation. The mechanisms leading to novel variation in newly formed allopolyploids may include gene dosage effects (resulting from more numerous copies of genome content), the reunion of divergent gene regulatory hierarchies, chromosomal rearrangements, and epigenetic remodeling, all of which affect gene content and/or expression levels. Many of these rapid changes may contribute to reproductive isolation and speciation. However, seed generated from interploidy crosses , such as between polyploids and their parent species, usually have aberrant endosperm development which impairs their viability, thus contributing to polyploid speciation . Polyploids may also interbreed with diploids and produce polyploid seeds, as observed in

8544-460: The production of aneuploid gametes. Natural or artificial selection for fertility can quickly stabilize meiosis in autopolyploids by restoring bivalent pairing during meiosis. Rapid adaptive evolution of the meiotic machinery, resulting in reduced levels of multivalents (and therefore stable autopolyploid meiosis) has been documented in Arabidopsis arenosa and Arabidopsis lyrata , with specific adaptive alleles of these species shared between only

8640-419: The production of polyploid cells. This process can be useful in plant breeding, especially when attempting to introgress germplasm across ploidal levels. Autopolyploids possess at least three homologous chromosome sets, which can lead to high rates of multivalent pairing during meiosis (particularly in recently formed autopolyploids, also known as neopolyploids) and an associated decrease in fertility due to

8736-562: The sauce from combining with the dough, and producing a crust that can be frozen and reheated without becoming rigid. There are frozen pizzas with raw ingredients and self-rising crusts. In the US, another form of pizza is available from take and bake pizzerias . This pizza is assembled in the store, then sold unbaked to customers to bake in their own ovens . Some grocery stores sell fresh dough along with sauce and basic ingredients, to assemble at home before baking in an oven. In restaurants, pizza can be baked in an oven with fire bricks above

8832-513: The word pizza comes from a notarial document written in Latin and dating to May 997 AD from Gaeta , demanding a payment of "twelve pizzas, a pork shoulder, and a pork kidney on Christmas Day, and 12 pizzas and a couple of chickens on Easter Day". Modern pizza evolved from similar flatbread dishes in Naples , Italy, in the 18th or early 19th century. Before that time, flatbread was often topped with ingredients such as garlic, salt, lard, and cheese. It

8928-440: The world (estimated in 2015 to be 14 million per year). As such, the city has been considered as one of the world capitals of pizza. The dish was introduced to Buenos Aires in the late 19th century with the massive Italian immigration , as part of a broader great European immigration wave to the country . Thus, around the same time that the iconic pizza Margherita was being invented in Italy, pizza were already being cooked in

9024-399: Was described as early as the 10th century by Ibn Wahshīya of Cairo . The North Africans called the product itrīya , from which Italian sources derived the term tria (or aletría in the case of Spanish sources) during the 15th century. Durum wheat ( Triticum turgidum ssp. durum ) is the 10th most cultivated cereal worldwide, with a total production of about 38 million tons. Most of

9120-412: Was induced in fish by Har Swarup (1956) using a cold-shock treatment of the eggs close to the time of fertilization, which produced triploid embryos that successfully matured. Cold or heat shock has also been shown to result in unreduced amphibian gametes, though this occurs more commonly in eggs than in sperm. John Gurdon (1958) transplanted intact nuclei from somatic cells to produce diploid eggs in

9216-540: Was then named after the queen, with an official letter of recognition from the queen's "head of service" remaining to this day on display in Esposito's shop, now called the Pizzeria Brandi. Later research cast doubt on this legend, undermining the authenticity of the letter of recognition, pointing that no media of the period reported about the supposed visit and that both the story and name Margherita were first promoted in

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