Tomato

The exact date of domestication is unknown; by 500 BC, it was already being cultivated in southern Mexico and probably other areas.^[10]: 13 The Pueblo people are thought to have believed that those who witnessed the ingestion of tomato seeds were blessed with powers of divination.^[11] The large, lumpy variety of tomato, a mutation from a smoother, smaller fruit, originated in Mesoamerica, and may be the direct ancestor of some modern cultivated tomatoes.^[10]: 15

The Aztecs raised several varieties of tomato, with red tomatoes called xitomatl and green tomatoes (physalis) called tomatl (

Spanish conquistador Hernán Cortés may have been the first to transfer a small yellow tomato to Europe after he captured the Aztec city of Tenochtitlan, now Mexico City, in 1521. The earliest discussion of the tomato in European literature appeared in a herbal written in 1544 by Pietro Andrea Mattioli, an Italian physician and botanist, who suggested that a new type of eggplant had been brought to Italy that was blood red or golden color when mature and could be divided into segments and eaten like an eggplant—that is, cooked and seasoned with salt, black pepper, and oil. It was not until ten years later that tomatoes were named in print by Mattioli as pomi d'oro, or "golden apples".^[10]: 13

After the Spanish colonization of the Americas, the Spanish distributed the tomato throughout their colonies in the Caribbean. They also took it to the Philippines, from where it spread to southeast Asia and then the entire Asian continent. The Spanish also brought the tomato to Europe. It grew easily in Mediterranean climates, and cultivation began in the 1540s. It was probably eaten shortly after it was introduced, and was certainly being used as food by the early 17th century in Spain.

China

The tomato was introduced to China, likely via the Philippines or Macau, in the 1500s. It was given the name 番茄 fānqié (foreign eggplant), as the Chinese named many foodstuffs introduced from abroad, but referring specifically to early introductions.^[15]

Italy

The recorded history of tomatoes in Italy dates back to at least 31 October 1548, when the house steward of Cosimo de' Medici, the grand duke of Tuscany, wrote to the Medici private secretary informing him that the basket of tomatoes sent from the grand duke's Florentine estate at Torre del Gallo "had arrived safely".^[16] Tomatoes were grown mainly as ornamentals early on after their arrival in Italy. For example, the Florentine aristocrat Giovanvettorio Soderini wrote how they "were to be sought only for their beauty", and were grown only in gardens or flower beds. The tomato's ability to mutate and create new and different varieties helped contribute to its success and spread throughout Italy. However, in areas where the climate supported growing tomatoes, their habit of growing to the ground suggested low status. They were not adopted as a staple of the peasant population because they were not as filling as other fruits already available. Additionally, both toxic and inedible varieties discouraged many people from attempting to consume or prepare any other varieties.^[17] In certain areas of Italy, such as Florence, the fruit was used solely as a tabletop decoration, until it was incorporated into the local cuisine in the late 17th or early 18th century.^[18] The earliest discovered cookbook with tomato recipes was published in Naples in 1692, though the author had apparently obtained these recipes from Spanish sources.^[10]: 17

Unique varieties were developed over the next several hundred years for uses such as dried tomatoes, sauce tomatoes, pizza tomatoes, and tomatoes for long-term storage. These varieties are usually known for their place of origin as much as by a variety name. For example, Pomodorino del Piennolo del Vesuvio is the "hanging tomato of Vesuvius", or the well known and highly prized San Marzano plum tomato grown in that region.^{[This paragraph needs citation(s)]}

Britain

Tomatoes were not grown in England until the 1590s. One of the earliest cultivators was John Gerard, a barber-surgeon. Gerard's Herbal, published in 1597, and largely plagiarized from continental sources, is also one of the earliest discussions of the tomato in England. Gerard knew the tomato was eaten in Spain and Italy. Nonetheless, he believed it was poisonous. (In fact, the plant and raw fruit do have low levels of tomatine, but are not generally dangerous. see below) Gerard's views were influential, and the tomato was considered unfit for eating (though not necessarily poisonous) for many years in Britain and its North American colonies.^[10]: 17

However, by the mid-18th century, tomatoes were widely eaten in Britain, and before the end of that century, the Encyclopædia Britannica stated the tomato was "in daily use" in soups, broths, and as a garnish. They were not part of the average person's diet, and though by 1820 they were described as "to be seen in great abundance in all our vegetable markets" and to be "used by all our best cooks", reference was made to their cultivation in gardens still "for the singularity of their appearance", while their use in cooking was associated with exotic Italian or Jewish cuisine.^[19] For example, in Elizabeth Blackwell's A Curious Herbal, it is described under the name "Love Apple (Amoris Pomum)" as being consumed with oil and vinegar in Italy, similar to consumption of cucumbers in the UK.^[20]

Middle East and North Africa

The tomato was introduced to cultivation in the Middle East by John Barker, British consul in Aleppo c. 1799 to 1825.^[21][22] Nineteenth century descriptions of its consumption are uniformly as an ingredient in a cooked dish. In 1881, it is described as only eaten in the region "within the last forty years".^[23] Today, the tomato is a crucial and ubiquitous part of Middle Eastern cuisine, served fresh in salads (e.g., Arab salad, Israeli salad, Shirazi salad and Turkish salad), grilled with kebabs and other dishes, made into sauces, and so on.^{[citation needed]}

United States

The earliest reference to tomatoes being grown in British North America is from 1710, when herbalist William Salmon reported seeing them in what is today South Carolina.^[10]: 25 They may have been introduced from the Caribbean. By the mid-18th century, they were cultivated on some Carolina plantations, and probably in other parts of the Southeast as well. Thomas Jefferson, who ate tomatoes in Paris, sent some seeds back to America.^[10]: 28 Some early American advocates of the culinary use of the tomato included Michele Felice Cornè and Robert Gibbon Johnson.^[24] Many Americans considered tomatoes to be poisonous at this time and, in general, they were grown more as ornamental plants than as food. In 1897, W. H. Garrison addressed the Medico-Legal Society of New York stating, "The belief was once transmitted that the tomato was sinisterly dangerous." He recalled in his youth tomatoes were dubbed "love-apples or wolf-apples" and they were shunned as "globes of the devil."^[25]

Alexander W. Livingston receives much credit for developing numerous varieties of tomato for both home and commercial gardeners.^[26]

Early tomato breeders included Henry Tilden in Iowa and a Dr. Hand in Baltimore.^[27] The U.S. Department of Agriculture's 1937 yearbook declared that "half of the major varieties were a result of the abilities of the Livingstons to evaluate and perpetuate superior material in the tomato." Livingston's first breed of tomato, the Paragon, was introduced in 1870, the beginning of a great tomato culture enterprise in the county. In 1875, he introduced the Acme, which was said to be involved in the parentage of most of the tomatoes introduced by him and his competitors for the next twenty-five years.^[28][29]

When Livingston began his attempts to develop the tomato as a commercial crop, his aim had been to grow tomatoes smooth in contour, uniform in size, and sweet in flavor. He eventually developed over seventeen different varieties of the tomato plant.^[28] Today, the crop is grown in every state in the Union.^[30]

Because of the long growing season needed for this heat-loving crop, several states in the US

The poor taste and lack of sugar in modern garden and commercial tomato varieties resulted from breeding tomatoes to ripen uniformly red. This change occurred after discovery of a mutant "u" phenotype in the mid-20th century, so named because the fruits ripened uniformly. This was widely cross-bred to produce red fruit without the typical green ring around the stem on uncross-bred varieties. Prior to general introduction of this trait, most tomatoes produced more sugar during ripening, and were sweeter and more flavorful.^[34][35]

Evidence has been found that 10–20% of the total carbon fixed in the fruit can be produced by photosynthesis in the developing fruit of the normal U phenotype. The u genetic mutation encodes a factor that produces defective chloroplasts with lower density in developing fruit, resulting in a lighter green colour of unripe fruit, and repression of sugars accumulation in the resulting ripe fruit by 10–15%. Perhaps more important than their role in photosynthesis, the fruit chloroplasts are remodelled during ripening into chlorophyll-free

Furthermore, breeders of modern tomato cultivars typically strive to produce tomato plants exhibiting improved yield, shelf life, size, and tolerance/resistance to various environmental pressures, including disease.[37]^[38] However, these breeding efforts have yielded unintended negative consequences on various tomato fruit attributes. For instance, linkage drag is a phenomenon that has been responsible for alterations in the metabolism of the tomato fruit. Linkage drag describes the introduction of an undesired trait or allele into a plant during backcrossing. This trait/allele is physically linked (or is very close) to the desired allele along the chromosome. In introducing the beneficial allele, there exists a high likelihood that the poor allele is also incorporated into the plant. Thus, breeding efforts attempting to enhance certain traits (for example: larger fruit size) have unintentionally altered production of chemicals associated with, for instance, nutritional value and flavor.^[37]

Breeders have turned to using wild tomato species as a source of alleles for the introduction of beneficial traits into modern tomato varieties. For example, wild tomato relatives may possess higher amounts of fruit solids (which are associated with greater sugar content) or resistance to diseases caused by microbes, such as resistance towards the early blight pathogen Alternaria solani. However, this tactic has limitations, for the incorporation of certain traits, such as pathogen resistance, can negatively impact other favorable phenotypes, such as fruit production.^[38][39]

Etymology

The word tomato comes from the Spanish tomate, which in turn comes from the Nahuatl word tomatl [ˈtomat͡ɬ] pronunciation^ⓘ, meaning 'swelling fruit';^[4] also 'fat water' or 'fat thing'.^[40] The native Mexican tomatillo is tomate. When Aztecs started to cultivate the fruit to be larger, sweeter and red, they called the new variety xitomatl (or jitomates) (pronounced [ʃiːˈtomatɬ]),^[2] ('plump with navel' or 'fat water with navel'). The specific name lycopersicum (from the 1753 book Species Plantarum) is of Greek origin (λύκοπερσικων, lykopersikon), meaning 'wolf peach'.^{[citation needed]}

Pronunciation

The usual pronunciations of tomato are /təˈmeɪtoʊ/ tə-MAY-toh (usual in North American English) and /təˈmɑːtoʊ/ tə-MAH-toh (usual in British English).^[41] The word's dual pronunciations were immortalized in Ira and George Gershwin's 1937 song "Let's Call the Whole Thing Off" ("You like /pəˈteɪtoʊ/ and I like /pəˈtɑːtoʊ/ / You like /təˈmeɪtoʊ/ and I like /təˈmɑːtoʊ/") and have become a symbol for nitpicking pronunciation disputes.^{[citation needed]} In this capacity, it has even become an American and British slang term: saying "/təˈmeɪtoʊ təˈmɑːtoʊ/" when presented with two choices can mean "What's the difference?" or "It's all the same to me".^{[citation needed]}

Botany

Description

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Tomato plants are vines, initially decumbent, typically growing 180 cm (6 ft) or more above the ground if supported, although erect bush varieties have been bred, generally 100 cm (3 ft 3 in) tall or shorter. Indeterminate types are "tender" perennials, dying annually in temperate climates (they are originally native to tropical highlands), although they can live up to three years in a greenhouse in some cases. Determinate types are annual in all climates.^{[citation needed]}

Tomato plants are

Tomato vines are typically pubescent, meaning covered with fine short hairs. The hairs facilitate the vining process, turning into roots wherever the plant is in contact with the ground and moisture, especially if the vine's connection to its original root has been damaged or severed.^{[citation needed]}

Most tomato plants have

The leaves are 10–25 cm (4–10 in) long, odd pinnate, with five to nine leaflets on petioles,^[44] each leaflet up to 8 cm (3 in) long, with a serrated margin; both the stem and leaves are densely glandular-hairy.^{[citation needed]}

Their

For propagation, the seeds need to come from a mature fruit, and must be lightly fermented to remove the gelatinous outer coating and then dried before use.[49]

Classification

In 1753,

Genetic evidence has now shown that Linnaeus was correct to put the tomato in the genus Solanum, making Solanum lycopersicum the correct name.^[1][52] Both names, however, will probably be found in the literature for some time. Two of the major reasons for considering the genera separate are the leaf structure (tomato leaves are markedly different from any other Solanum), and the biochemistry (many of the alkaloids common to other Solanum species are conspicuously absent from the tomato). On the other hand, hybrids of tomato and diploid potato can be created in the lab by somatic fusion, and are partially fertile,^[53] providing evidence of the close relationship between these species.

Genetics and genetic modification

Genome

An international consortium of researchers from 10 countries, began sequencing the tomato genome in 2004.^[54][55] A prerelease version of the genome was made available in December 2009.^[56] The complete genome for the cultivar Heinz 1706 was published on 31 May 2012 in Nature.^[57] The latest reference genome published in 2021 had 799 MB and encodes 34,384 (predicted) proteins, spread over 12 chromosomes.^[58]

Genetic modification

Since many other fruits, like

The first commercially available genetically modified food was a tomato called Flavr Savr, which was engineered to have a longer shelf life.^[60] However, it is no longer commercially available. Scientists are continuing to develop tomatoes with new traits not found in natural crops, such as increased resistance to pests or environmental stresses or better flavor.^{[citation needed]}

Breeding

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The Tomato Genetic Resource Center,

Tomato cultivars vary widely in their resistance to disease. Modern hybrids focus on improving disease resistance over the heirloom plants.

Various forms of mildew and blight are common tomato afflictions, which is why tomato cultivars are often marked with a combination of letters that refer to specific disease resistance. The most common letters are:

• LB – '
  late blight'^[67]
• V – verticillium wilt
• F – fusarium wilt strain I
• FF – fusarium wilt strain I and II
• N – nematodes
• T – tobacco mosaic virus, and
• A – alternaria.

A common tomato disease is tobacco mosaic virus. Handling cigarettes and other infected tobacco products can transmit the virus to tomato plants.^[68]

Another particularly dreaded disease is curly top, carried by the beet leafhopper, which interrupts the lifecycle. As the name implies, it has the symptom of making the top leaves of the plant wrinkle up and grow abnormally.^{[citation needed]}

After an insect attack tomato plants produce systemin, a plant peptide hormone. Systemin activates defensive mechanisms, such as the production of protease inhibitors to slow the growth of insects. The hormone was first identified in tomatoes, but similar proteins have been identified in other species since.^[72]

Other disorders

Although not a disease as such, irregular supplies of water can cause growing or ripening fruit to split. Besides cosmetic damage, the splits may allow decay to start, although growing fruits have some ability to heal after a split. In addition, a deformity called cat-facing can be caused by pests, temperature stress, or poor soil conditions. Affected fruit usually remains edible, but its appearance may be unsightly.

Companion plants

Tomatoes serve, or are served by, a large variety of

The devastating tomato hornworm has a major predator in various

Basil is popularly recommended as a companion plant to the tomato. Common claims are that basil may deter pests or improve tomato flavor. However, in double-blind taste tests, basil did not significantly affect the taste of tomatoes when planted adjacent to them.^[75][76]

Tomato plants can protect asparagus from asparagus beetles, because they contain solanine that kills this pest,^{[citation needed]} while asparagus plants contain asparagusic acid that repels nematodes known to attack tomato plants.^[77] Marigolds also repel nematodes.^[78][79][80]

Pollination

In the wild, original state, tomatoes required cross-

As tomatoes were moved from their native areas, their traditional pollinators (probably a species of halictid bee) did not move with them.^[81] The trait of self-fertility became an advantage, and domestic cultivars of tomato have been selected to maximize this trait.^[81]

This is not the same as

Pollination and fruit formation depend on meiosis. Meiosis is central to the processes by which diploid microspore mother cells within the anther give rise to haploid pollen grains, and megaspore mother cells in ovules that are contained within the ovary give rise to haploid nuclei. Union of haploid nuclei from pollen and ovule (fertilization) can occur either by self- or cross-pollination. Fertilization leads to the formation of a diploid zygote that can then develop into an embryo within the emerging seed. Repeated fertilizations within the ovary are accompanied by maturation of the ovary to form the tomato fruit.

Homologs of the

Tomatoes are often grown in

Greenhouse tomato production in large-acreage commercial greenhouses and owner-operator stand-alone or multiple-bay greenhouses is on the increase, providing fruit during those times of the year when field-grown fruit is not readily available. Smaller sized fruit (cherry and grape), or cluster tomatoes (fruit-on-the-vine) are the fruit of choice for the large commercial greenhouse operators while the beefsteak varieties are the choice of owner-operator growers.^[84]

Picking and ripening

To facilitate transportation and storage, tomatoes are often picked unripe (green) and ripened in storage with

A machine-harvestable variety of tomato (the "square tomato") was developed in the 1950s by University of California, Davis's Gordie C. Hanna, which, in combination with the development of a suitable harvester, revolutionized the tomato-growing industry. This type of tomato is grown commercially near plants that process and can tomatoes, tomato sauce, and tomato paste. They are harvested when ripe and are flavorful when picked. They are harvested 24 hours a day, seven days a week during a 12- to 14-week season, and immediately transported to packing plants, which operate on the same schedule. California is a center of this sort of commercial tomato production and produces about a third of the processed tomatoes produced in the world.^[86]

In 1994,

Several US states have adopted the tomato as a state fruit or vegetable (see above). Tomatoes have been designated the state vegetable of New Jersey. Arkansas took both sides by declaring the South Arkansas Vine Ripe Pink Tomato both the state fruit and the state vegetable in the same law, citing both its culinary and botanical classifications. In 2009, the state of Ohio passed a law making the tomato the state's official fruit. Tomato juice has been the official beverage of Ohio since 1965. Alexander W. Livingston, of Reynoldsburg, Ohio, played a large part in popularizing the tomato in the late 19th century; his efforts are commemorated in Reynoldsburg with an annual Tomato Festival.

Flavr Savr was the first commercially grown genetically engineered food licensed for human consumption.^[117]

Tomatoes are a popular "nonlethal" throwing weapon in mass protests, and there was a common tradition of throwing rotten tomatoes at bad performers on a stage during the 19th century; today this is usually referenced as a metaphor. Embracing it for this protest connotation, the Dutch Socialist party adopted the tomato as their logo.

The US city of Reynoldsburg, Ohio calls itself "The Birthplace of the Tomato", claiming the first commercial variety of tomato was bred there in the 19th century.^[28]

"Rotten Tomatoes" is an American review-aggregation website for film and television. The name "Rotten Tomatoes" derives from the practice of audiences throwing rotten tomatoes when disapproving of a poor stage performance.

Gallery

• 
  Various heirloom tomato (heritage tomato in British English) cultivars
• 
  Homegrown multicolored tomatoes
• 
  A variety of cultivars, including

  Black Krim (lower left) and Green Zebra

  (top left)
• 
  Red tomatoes with PLU code in a supermarket

See also

References

Further reading

• David Gentilcore. Pomodoro! A History of the Tomato in Italy (Columbia University Press, 2010), scholarly history
• Tieman, D; Bliss, P; McIntyre, LM; Blandon-Ubeda, A; Bies, D; Odabasi, AZ; Rodríguez, GR; van der Knaap, E; Taylor, MG; Goulet, C; Mageroy, MH; Snyder, DJ; Colquhoun, T; Moskowitz, H; Clark, DG; Sims, C; Bartoshuk, L; Klee, HJ (5 June 2012). "The Chemical Interactions Underlying Tomato Flavor Preferences". Current Biology. 22 (11): 1035–1039.
  PMID 22633806
  .

External links

Wikibooks Cookbook has a recipe/module on

• Tomato

Wikibooks has a book on the topic of: Horticulture

Wikibooks has a book on the topic of: Tomato

Wikispecies has information related to Solanum lycopersicum.

Wikimedia Commons has media related to Solanum lycopersicum and Tomatoes.

Look up tomato in Wiktionary, the free dictionary.

• Tomato Genome Sequencing Project – Sequencing of the twelve tomato chromosomes.
• Tomato core collection database – Phenotypes and images of 7,000 tomato cultivars