{{Short description|Edible berry}} {{other uses}} {{good article}} {{pp-move}} {{protection padlock |small=yes}} {{Use dmy dates |date=August 2023}} {{Speciesbox | name = Tomato | image = Tomato je.jpg | image_caption = | genus = Solanum | species = lycopersicum | authority = L. | synonyms = {{ubl |''Lycopersicon lycopersicum'' {{small |(L.H. Karst.)}} |''Lycopersicon esculentum'' {{small |(Mill.)}}}} |synonyms_ref = <ref name="NHM">{{cite web |quote=Molecular phylogenetic analyses have established that the formerly segregate genera ''Lycopersicon'', ''Cyphomandra'', ''Normania'', and ''Triguera'' are nested within ''Solanum'', and all species of these four genera have been transferred to ''Solanum'' |url=http://solanaceaesource.org/content/phylogeny-0 |title=Phylogeny}}</ref> }}
The '''tomato''' ({{IPAc-en|US|t|ə|ˈ|m|eɪ|t|oʊ}}, {{IPAc-en|UK|t|ə|ˈ|m|aː|t|oʊ}}; '''''Solanum lycopersicum''''') is a plant whose fruit is an edible berry that is eaten as a vegetable. The tomato is a member of the nightshade family that includes tobacco, potato, and chili peppers. It originated from western South America, and may have been domesticated there, in Mexico, or in Central America. The Spanish introduced tomatoes to Eurasia in the Columbian exchange in the 16th century.
Tomato plants are vines, largely annual, and vulnerable to frost, though sometimes living longer in greenhouses. The flowers are able to self-fertilize. Modern varieties have been bred to ripen uniformly red, in a process that has impaired the fruit's sweetness and flavor. There are thousands of cultivars, varying in size, color, shape, and flavor. Tomatoes are attacked by many insect pests and nematodes and are subject to diseases caused by viruses, mildew, and blight fungi.
The tomato has a strong savory umami flavor and is an important ingredient in cuisines around the world. Tomatoes are widely used in sauces for pasta and pizza, in soups such as gazpacho and tomato soup, in salads and condiments like salsa and ketchup, and in various curries. Tomatoes are also consumed as juice and in beverages such as the Bloody Mary cocktail.
== Naming ==
=== Etymology ===
The word ''tomato'' comes from the Spanish {{lang|es |tomate}}, which in turn comes from the Nahuatl word {{lang|nci |tomatl}} {{IPA|nah |ˈtomat͡ɬ |}} {{Pronunciation |Tomatl.ogg}}.<ref>{{OEtymD |tomato |accessdate=2024-07-13}}</ref> The specific name ''lycopersicum'', meaning 'wolf peach', originated with Galen, who used it to denote a plant that has never been identified. Luigi Anguillara speculated in the 16th century that Galen's ''lycopersicum'' might be the tomato, and despite the impossibility of this identification, ''lycopersicum'' entered scientific use as a name for the fruit.<ref>{{cite journal |last=Sabine |first=Joseph |date=1820 |title=On the Love Apple or Tomato |url=https://books.google.com/books?id=7ALkF3ZIfCkC&pg=PA343 |journal=Transactions of the Horticultural Society of London |volume=3 |page=343 f}}</ref>
=== Pronunciation ===
The usual pronunciations of ''tomato'' are {{IPAc-en|t|ə|ˈ|m|eɪ|t|oʊ}} (in North American English) and {{IPAc-en|t|ə|ˈ|m|ɑː|t|oʊ}} (in British English).<ref name="Cambridge Dictionaries Online-2015">{{cite web |url=http://dictionary.cambridge.org/dictionary/british/tomato |title=English definition of 'tomato' |author=<!--Staff writer(s); no by-line.--> |year=2015 |publisher=Cambridge University Press |access-date=15 May 2015}}</ref> The word's dual pronunciations were immortalized in Ira and George Gershwin's 1937 song "Let's Call the Whole Thing Off" ("You like {{IPAc-en|p|ə|ˈ|t|eɪ|t|oʊ}} and I like {{IPAc-en|p|ə|ˈ|t|ɑː|t|oʊ}} / You like {{IPAc-en|t|ə|ˈ|m|eɪ|t|oʊ}} and I like {{IPAc-en|t|ə|ˈ|m|ɑː|t|oʊ}}").<ref>{{cite web |title=Let's Call The Whole Thing Off: Song by Ella Fitzgerald |url=https://www.google.co.uk/search?q=let%27s+call+the+whole+thing+off+lyrics&sca_esv=3145f941300c1a56&source=hp&ei=QGcOZ6SnJKaFhbIPusWC6Qw&iflsig=AL9hbdgAAAAAZw51UH_jTwdasztauAPXIIQTt3t9qyff&oq=%22let%27s+call+%22&gs_lp=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&sclient=gws-wiz |website=Google |access-date=15 October 2024}}</ref>
== History ==
[[File:Naturalis Biodiversity Center - Solanum lycopersicum var. lycopersicum - old tomato herbarium sheet.jpg|thumb |upright |''Solanum lycopersicum'' var. ''lycopersicum'': one of the oldest surviving tomato specimens. Page from the ''En Tibi Herbarium'', 1558. Naturalis Leiden.]]
The likely wild ancestor of the tomato, the red-fruited ''Solanum pimpinellifolium'', is native to western South America, where it was probably first domesticated. The resulting domesticated plant, ancestral to the modern large-fruited tomato varieties, was probably the cherry tomato, ''S. lycopersicum'' var. ''cerasiforme''.<ref name="Lin-2014">{{cite journal |last1=Lin |first1=Tao |last2=Zhu |first2=Guangtao |last3=Zhang |first3=Junhong |last4=Xu |first4=Xiangyang |last5=Yu |first5=Qinghui |last6=Zheng |first6=Zheng |last7=Zhang |first7=Zhonghua |last8=Lun |first8=Yaoyao |last9=Li |first9=Shuai |last10=Wang |first10=Xiaoxuan |last11=Huang |first11=Zejun |last12=Li |first12=Junming |last13=Zhang |first13=Chunzhi |last14=Wang |first14=Taotao |last15=Zhang |first15=Yuyang |last16=Wang |first16=Aoxue |last17=Zhang |first17=Yancong |last18=Lin |first18=Kui |last19=Li |first19=Chuanyou |last20=Xiong |first20=Guosheng |last21=Xue |first21=Yongbiao |last22=Mazzucato |first22=Andrea |last23=Causse |first23=Mathilde |last24=Fei |first24=Zhangjun |last25=Giovannoni |first25=James J. |last26=Chetelat |first26=Roger T. |last27=Zamir |first27=Dani |last28=Städler |first28=Thomas |last29=Li |first29=Jingfu |last30=Ye |first30=Zhibiao |last31=Du |first31=Yongchen |last32=Huang |first32=Sanwen |display-authors=5 |title=Genomic analyses provide insights into the history of tomato breeding |journal=Nature Genetics |volume=46 |issue=11 |date=12 October 2014 |doi=10.1038/ng.3117 |pages=1220–1226|pmid=25305757 }}</ref><ref name="Estabrook-2015">{{cite magazine |last=Estabrook |first=Barry |title=Why Is This Wild, Pea-Sized Tomato So Important? |url=https://www.smithsonianmag.com/travel/why-wild-tiny-pimp-tomato-so-important-180955911 |date=22 July 2015 |magazine=Smithsonian Journeys Quarterly |access-date=13 January 2020 |archive-date=13 January 2020 |archive-url=https://web.archive.org/web/20200113211431/https://www.smithsonianmag.com/travel/why-wild-tiny-pimp-tomato-so-important-180955911/ |url-status=live }}</ref> However, genomic analysis suggests that the domestication process may have been more complex than this. ''S. lycopersicum'' var. ''cerasiforme'' may have existed before domestication, while traits supposedly typical of domestication may have been reduced in that variety and then reselected (in a case of convergent evolution) in the cultivated tomato. The analysis predicts that var. ''cerasiforme'' appeared around 78,000 years ago, while the cultivated tomato originated around 7,000 years ago (5,000 BCE), with substantial uncertainty, making it unclear how humans may have been involved in the process.<ref name="Razifard-2020">{{cite journal |last=Razifard |first=Hamid |display-authors=etal |title=Genomic evidence for complex domestication history of the cultivated tomato in Latin America |journal=Molecular Biology and Evolution |volume=37 |issue=4 |year=2020 |pages=1118–1132 |doi=10.1093/molbev/msz297 |pmid=31912142 |pmc=7086179 |url=https://academic.oup.com/mbe/article-pdf/37/4/1118/32960114/msz297.pdf}}</ref>
The Spanish first introduced tomatoes to Europe, where they became used in Spanish food. Elsewhere in Europe, its first use was ornamental, not least because it was understood to be related to the nightshades and assumed to be poisonous.<ref name="Britannica-2018">{{cite web |url=https://www.britannica.com/plant/tomato |title=Tomato |publisher=Encyclopaedia Britannica |date=4 January 2018 |access-date=15 January 2018 |archive-date=20 August 2023 |archive-url=https://web.archive.org/web/20230820082833/https://www.britannica.com/plant/tomato |url-status=live }}</ref>
=== Mesoamerica ===
While the tomato's wild ancestors grow in Chile, Peru, Ecuador, the fruit was domesticated by precontact Central American farmers.{{sfn|Smith|1994|p=15}} The exact date of domestication is unknown; by 500 BCE, it was already being cultivated in southern Mexico and probably other areas.{{sfn|Smith|1994|p=13}} A 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.{{sfn|Smith|1994|p=15}}
The Aztecs raised several varieties of tomato, with red tomatoes called {{lang|nci|xitomatl}}.<ref>{{cite book |last=Townsend |first=Richard F. |title=The Aztecs |publisher=Thames and Hudson |year=2000 |pages=180–181}}</ref> Bernardino de Sahagún reported seeing a great variety of tomatoes in the Aztec market at Tenochtitlán (Mexico City): "large tomatoes, small tomatoes, leaf tomatoes, sweet tomatoes, large serpent tomatoes, nipple-shaped tomatoes", and tomatoes of all colors from the brightest red to the deepest yellow.<ref>{{cite book |last=Silvertown |first=J. |year=2017 |title=Vegetables—Variety. Dinner with Darwin: Food, drink, and evolution |publisher=University of Chicago Press |page=102}}</ref> Sahagún mentioned Aztecs cooking various sauces, some with tomatoes of different sizes, serving them in city markets: "foods sauces, hot sauces; ... with tomatoes, ... sauce of large tomatoes, sauce of ordinary tomatoes, ..."<ref name="Coe-2015">{{cite book |last=Coe |first=Sophie D. |author-link=Sophie Coe |title=America's First Cuisines |publisher=University of Texas Press |location=Austin, Texas |date=2015 |orig-year=1994 |pages=108–118 [117] |url=https://books.google.com/books?id=hKVbCgAAQBAJ&pg=PA117 |isbn=978-1477309711}}</ref>
=== Spanish distribution === {{further|Columbian exchange}} [[File:Arrival of Cortés in Veracruz and Reception by Moctezuma's Ambassadors Painting.jpg|thumb|Soon after Hernán Cortés's arrival in Mexico (portrayed here) and his conquest of the Aztecs, the tomato was brought to Europe in the Columbian exchange.<ref name="López-Terrada"/>]]
The Spanish conquistador Hernán Cortés's capture of Tenochtitlan in 1521 initiated the widespread cultural and biological interchange called the Columbian exchange. The tomato was cultivated in Europe only a few years after that event,<ref name="López-Terrada"/> by the 1540s, and grew easily in the Mediterranean climates. The earliest mention of the tomato in European literature appeared in Pietro Andrea Mattioli's 1544 herbal. He suggested that a new type of eggplant had been brought to Italy. He stated that it 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. Ten years later, Mattioli named the fruits in print as {{lang|it|pomi d'oro}}, or "golden apples".{{sfn|Smith|1994|p=13}}
It was probably eaten shortly after it was introduced, and tomatoes were used as food by the early 17th century in Spain, as documented in the 1618 play ''La octava maravilla'' by Lope de Vega with "lovelier than ... a tomato in season".<ref name="López-Terrada">{{cite web |last=López-Terrada |first=Maríaluz |title=The History of the Arrival of the Tomato in Europe: An Initial Overview |url=https://traditom.eu/fileadmin/traditom/downloads/TRADITOM_History_of_the_arrival_of_the_tomato_in_Europe.pdf |website=Traditom |access-date=9 October 2024 |archive-date=9 October 2024 |archive-url=https://web.archive.org/web/20241009110136/https://traditom.eu/fileadmin/traditom/downloads/TRADITOM_History_of_the_arrival_of_the_tomato_in_Europe.pdf |url-status=dead }}</ref>
After the Spanish colonization of the Americas, the Spanish distributed the tomato throughout their colonies in the Caribbean. After they introduced it to the Philippines, it spread to Southeast Asia and then across Asia.<ref name="Hancock-2022">{{cite book |last=Hancock |first=James F. |title=World Agriculture Before and After 1492 |chapter=Dispersal of New World Crops into the Old World |publisher=Springer International Publishing |publication-place=Cham |year=2022 |isbn=978-3-031-15522-2 |doi=10.1007/978-3-031-15523-9_9 |pages=111–133}}</ref>
=== China ===
The tomato was introduced to China, likely via the Philippines or Macau, in the 16th century. It was given the name 番茄 {{transliteration |zh |fānqié}} (foreign eggplant), as the Chinese named many foodstuffs introduced from abroad, but referring specifically to early introductions.<ref>{{cite book |last1=Kiple |first1=Kenneth F. |last2=Ornelas |first2=Kriemhild Coneè |title=The Cambridge World History of Food |volume=1 |url=https://books.google.com/books?id=RSSkDNzKQacC |year=2000 |publisher=Cambridge University Press |isbn=978-0-521-40214-9 |page=357}}</ref>
=== Italy === [[File:TomateSanMarzano.jpg|thumb |upright |The San Marzano is a well-known plum tomato highly prized for making pizza.]] In 1548, 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".<ref>{{cite web |url=https://pomodoro.museidelcibo.it/en/audio-guides-2/ |title=Tomato Museum: 05 – The History of Tomato |publisher=I Musei del Cibo della provincia di Parma |access-date=30 June 2022 |archive-date=19 June 2022 |archive-url=https://web.archive.org/web/20220619212846/https://pomodoro.museidelcibo.it/en/audio-guides-2/ |url-status=live }}</ref> 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 close 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 crops. Additionally, both toxic and inedible varieties discouraged many people from attempting to consume or prepare any other varieties.<ref>{{cite book |last=Gentilcore |first=David |year=2010 |title=Pomodoro! A History of the Tomato in Italy |publisher=Columbia University Press |isbn=978-0-231-15206-8}}.</ref> 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.<ref>{{cite book |last1=Staller |first1=John |last2=Carrasco |first2=Michael |title=Pre-Columbian Foodways: Interdisciplinary Approaches to Food, Culture, and Markets in Ancient Mesoamerica |date=2009 |publisher=Springer Science & Business Media |isbn=978-144190471-3 |page=44 |url=https://books.google.com/books?id=FJrr9i6HRp0C&q=tabletop+decoration |access-date=3 May 2021}}</ref> The earliest discovered cookbook with tomato recipes was published in Naples in 1692, though the author had apparently obtained these recipes from Spanish sources.{{sfn|Smith|1994|p=17}}
Varieties were developed over the following centuries for drying, for sauce, for pizzas, and for long-term storage. These varieties are usually known for their place of origin as much as by a variety name. For example, there is the {{lang|it|Pomodorino del Piennolo del Vesuvio}}, the "hanging tomato of Vesuvius", and the well known and highly prized San Marzano tomato grown in that region, with a European protected designation of origin certification.<ref>{{cite journal |last1=Rao |first1=R. |last2=Corrado |first2=G. |last3=Bianchi |first3=M. |last4=Di Mauro |first4=A. |title=(GATA)4 DNA fingerprinting identifies morphologically characterized 'San Marzano' tomato plants |journal=Plant Breeding |date=21 March 2006 |volume=125 |issue=2 |pages=173–176 |doi=10.1111/j.1439-0523.2006.01183.x |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1439-0523.2006.01183.x |access-date=13 April 2023|url-access=subscription }}</ref>
=== Britain ===
thumb|Tomatoes for sale in a UK supermarket
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. Gerard's views were influential, and the tomato was considered unfit for eating for many years in Britain and its North American colonies.{{sfn|Smith|1994|p=17}} By 1820, tomatoes 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.<ref>{{cite news |title=Love-apple, or Tomato Berry. – Love apples are now to be seen in great abundance at all our vegetable markets |newspaper=The Times |date=22 September 1820 |page=3}}</ref> For example, in Elizabeth Blackwell's ''A Curious Herbal'', it is described under the name "Love Apple ({{lang|la|Amoris Pomum}})" as being consumed with oil and vinegar in Italy, similar to consumption of cucumbers in the UK.<ref>{{cite book |last=Blackwell |first=Elizabeth |author-link=Elizabeth Blackwell (illustrator) |url=https://collections.nlm.nih.gov/ext/dw/2449056RX1/PDF/2449056RX1.pdf |url-status=dead |archive-date=6 June 2023 |archive-url=https://web.archive.org/web/20230606092105/https://collections.nlm.nih.gov/ext/dw/2449056RX1/PDF/2449056RX1.pdf |page=342 (plate 133) |title=A curious herbal: containing five hundred cuts, of the most useful plants, which are now used in the practice of physick: engraved on folio copper plates, after drawings taken from life |year=1737}}</ref><!--this image should be here in the article--> In 1963, ''The New York Times'' gave an explanation of the name 'Love Apple' as a French misreading of the Italian {{lang|it|pomo dei Mori}} ("the Moors' apple") as {{lang|fr|pomme d'amour}}, ("apple of love").<ref>{{cite news |last=Alma |first=C.M. |date=7 April 1963 |title=Aztecs' Tomatl is the Modern Tomato: From Spain to Italy on a Bush |newspaper=The New York Times |url=https://www.proquest.com/docview/116565156|id={{ProQuest|116565156}} }}</ref>
=== Middle East ===
The tomato was introduced to cultivation in the Middle East by John Barker, British consul in Aleppo {{circa |1799 to 1825}}.<ref>{{cite journal |last=Bergougnoux |first=Véronique |title=The history of tomato: from domestication to biopharming |journal=Biotechnology Advances |volume=32 |issue=1 |year=2014 |pages=170–189 |doi=10.1016/j.biotechadv.2013.11.003 |pmid=24211472 |url=https://www.academia.edu/80426432}}</ref><ref>{{cite web |url=http://yourarchives.nationalarchives.gov.uk/index.php?title=British_Consuls_in_Aleppo |title=British Consuls in Aleppo |publisher=Yourarchives.nationalarchives.gov.uk |date=26 January 2009 |access-date=2 April 2009 |archive-date=8 September 2011 |archive-url=https://web.archive.org/web/20110908042910/http://yourarchives.nationalarchives.gov.uk/index.php?title=British_Consuls_in_Aleppo |url-status=live }}</ref><ref>{{cite book |chapter-url=https://books.google.com/books?id=6QUZAAAAYAAJ&pg=PA519 |chapter=Syria under the last five Turkish Sultans |title=Appletons' Journal |volume=1 |publisher=D. Appleton and Co. |year=1876 |page=519 }}</ref> 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".<ref>{{cite journal |url=https://books.google.com/books?id=pnkrAAAAYAAJ&pg=PA223 |title=Natural History, Science, &c |journal=The Friend |volume=54 |year=1881 |page=223}}</ref>
=== United States ===
[[File:NRCSHI07018 - Hawaii (716072)(NRCS Photo Gallery).jpg|thumb|Plum, cherry, and grape varieties in Hawaii.]]
The earliest reference to tomatoes being grown in British North America is from 1710, when herbalist William Salmon saw them in what is today South Carolina,{{sfn|Smith|1994|p=25}} perhaps introduced from the Caribbean. By the mid-18th century, they were cultivated on some Carolina plantations, and probably in other parts of the Southeast. Thomas Jefferson, who ate tomatoes in Paris, sent some seeds back to America.{{sfn|Smith|1994|p=28}} Some early American advocates of the culinary use of the tomato included Michele Felice Cornè and Robert Gibbon Johnson.<ref>{{cite journal |last=McCue |first=George Allen |title=The History of the Use of the Tomato: An Annotated Bibliography |journal=Annals of the Missouri Botanical Garden |volume=39 |issue=4 |date=November 1952 |publisher=Missouri Botanical Garden Press |doi=10.2307/2399094 |pages=336–338 |jstor=2399094 |bibcode=1952AnMBG..39..289M |url=https://www.biodiversitylibrary.org/part/22432 |archive-date=8 April 2022 |access-date=19 November 2021 |archive-url=https://web.archive.org/web/20220408131134/https://www.biodiversitylibrary.org/part/22432 |url-status=live }}</ref> 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 stated, "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 shunned as "globes of the devil".<ref>{{cite book |last1=Harrald |first1=Chris |last2=Watkins |first2=Fletcher |title=The cigarette book: the history and culture of smoking |publisher=Skyhorse Publishing |year=2010 |page=185}}</ref>
When Alexander W. Livingston (1821–1898) began developing 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 varieties.{{sfn|Smith|1994|p=152}}<ref>{{cite web |url=https://www.agmrc.org/commodities-products/vegetables/tomatoes/ |title=Tomatoes |website=AgMRC |date=March 2017 |access-date=25 May 2018}}</ref> 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. In 1875, he introduced the Acme, said to be in the parentage of most cultivars for the next twenty-five years. Other early breeders included Henry Tilden in Iowa and a Dr. Hand in Baltimore.<ref name="Boswell-1937">Boswell, Victor R. "[https://archive.org/details/yoa1937 Improvement and Genetics of Tomatoes, Peppers, and Eggplant]", ''Yearbook of Agriculture, 1937,'' p. 179. U.S. Department of Agriculture. Accessed 25 May 2018/</ref>
Because of the tomato's need for heat and a long growing season, several states in the Sun Belt became major producers, particularly Florida and California. In California, tomatoes are grown under irrigation for both the fresh market and for canning and processing. The University of California, Davis's C.M. Rick Tomato Genetics Resource Center maintains a gene bank of <!--tomato -->wild relatives, monogenic mutants and genetic stocks.<ref>{{cite web |url=http://tgrc.ucdavis.edu/ |title=C.M. Rick Tomato Genetics Resource Center |publisher=University of California, Davis |access-date=2 April 2009 |archive-date=14 May 2011 |archive-url=https://web.archive.org/web/20110514051356/http://tgrc.ucdavis.edu/ |url-status=live }}</ref> Research on processing tomatoes is also conducted by the California Tomato Research Institute in Escalon, California.<ref>[https://web.archive.org/web/20100709222618/http://www.tomatonet.org/ctri.htm California Tomato Research Institute]. tomatonet.org</ref> In California, growers have used a method of cultivation called dry-farming, especially with Early Girl tomatoes. This technique encourages the plant to send roots deep to find existing moisture.<ref name="Kornei-2023">{{cite web |last=Kornei |first=Katherine |title=Dry farming could help agriculture in the western U.S. amid climate change |url=https://www.sciencenews.org/article/dry-farming-agriculture-climate-change |website=Science News |access-date=9 October 2024 |date=9 March 2023 |archive-date=13 January 2025 |archive-url=https://web.archive.org/web/20250113142526/https://www.sciencenews.org/article/dry-farming-agriculture-climate-change |url-status=live }}</ref>
== Botany ==
=== Description ===
Tomato plants are vines, becoming decumbent, and can grow up to {{cvt|3|m}}; bush varieties are generally no more than {{cvt|100|cm|ftin|0}} tall. They are tender perennials, often grown as annuals.<ref name="MissouriBG">{{cite web |title=Solanum lycopersicum |url=http://www.missouribotanicalgarden.org/PlantFinder/PlantFinderDetails.aspx?taxonid=454685 |publisher=Missouri Botanical Garden |access-date=22 October 2024 |archive-date=1 December 2024 |archive-url=https://web.archive.org/web/20241201193332/http://www.missouribotanicalgarden.org/PlantFinder/PlantFinderDetails.aspx?taxonid=454685 |url-status=live }}</ref><ref name="SEINet">{{cite web |title=Solanum lycopersicon L. |url=https://swbiodiversity.org/seinet/taxa/index.php?taxon=Solanum%20lycopersicum |website=SEINet |access-date=22 October 2024 |archive-date=1 December 2024 |archive-url=https://web.archive.org/web/20241201190312/https://swbiodiversity.org/seinet/taxa/index.php?taxon=Solanum%20lycopersicum |url-status=live }}</ref>
Tomato plants are dicots. They grow as a series of branching stems, with a terminal bud at the tip that does the actual growing. When the tip eventually stops growing, whether because of pruning or flowering, lateral buds take over and grow into new, fully functional, vines.<ref>{{cite web |url=http://www.ncsu.edu/sustainable/profiles/bot_tom.html |archive-url=https://web.archive.org/web/20091126050832/http://www.ncsu.edu/sustainable/profiles/bot_tom.html |archive-date=26 November 2009 |title=Crop Profiles – Tomato |access-date=27 October 2008 |first=M. |last=Peet}}</ref> 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.<ref name="SingaporeNP"/> The leaves are {{cvt|10|–|25|cm|in|0|abbr=on}} long, odd pinnate, with five to nine leaflets on petioles, each leaflet up to {{cvt|8|cm|in|0|abbr=on}} long, with a serrated margin; both the stem and leaves are densely glandular-hairy.<ref name="North Carolina Extension Gardener Plant Toolbox">{{cite web |title=Solanum lycopersicum |url=https://plants.ces.ncsu.edu/plants/solanum-lycopersicum/ |publisher=North Carolina Extension Gardener Plant Toolbox |access-date=9 October 2024}}</ref>
Tomato flowers are bisexual and are able to self fertilize. As tomatoes were moved from their native areas, their traditional pollinators (probably a species of halictid bee) did not move with them.<ref name="Sharma-2012">{{cite book |last=Sharma |first=V.P. |title=Nature at Work – the Ongoing Saga of Evolution |url=https://books.google.com/books?id=o4qwjov434kC&pg=PA41 |year=2012 |publisher=Springer |isbn=978-81-8489-991-7 |page=41}}</ref> The trait of self-fertility became an advantage, and domestic cultivars of tomato have been selected to maximize this trait.<ref name="Sharma-2012"/> This is not the same as self-pollination, despite the common claim that tomatoes do so. That tomatoes pollinate themselves poorly without outside aid is clearly shown in greenhouse situations, where pollination must be aided by artificial wind, vibration of the plants, or by cultured bumblebees.<ref>{{cite book |last1=Frankie |first1=Gordon |last2=Thorp |first2=Robbin |last3=Coville |first3=Rollin |last4=barbara |first4=Ertter |last5=California Native Plant Society |title=California bees & blooms: a guide for gardeners and naturalists |date=2014 |publisher=Heydey |location=Berkeley, CA |isbn=9781597142946}}</ref>
The flowers develop on the apical meristem. They have the anthers fused along the edges, which form a column surrounding the pistil's style. The anthers bend into a cone-like structure, surrounding the stigma. The flowers are {{cvt|1|–|2|cm|in|1|abbr=on}} across, yellow, with five pointed lobes on the corolla; they are borne in a cyme of three to twelve together.<ref name="SingaporeNP">{{Cite web |title=Solanum lycopersicum |url=https://www.nparks.gov.sg/florafaunaweb/flora/5/7/5772 |access-date=9 October 2024 |publisher=Singapore National Parks |archive-date=9 October 2024 |archive-url=https://web.archive.org/web/20241009193330/https://www.nparks.gov.sg/florafaunaweb/flora/5/7/5772 |url-status=live }}</ref><ref name="RHS">{{cite web |title=Solanum lycopersicum: tomato |url=https://www.rhs.org.uk/plants/293619/solanum-lycopersicum/details |publisher=Royal Horticultural Society |access-date=9 October 2024}}</ref>
The fruit develops from the ovary of the plant after fertilization, its flesh comprising the pericarp walls. The fruit contains locules, hollow spaces full of seeds. These vary among cultivated varieties. Some smaller varieties have two locules; globe-shaped varieties typically have three to five; beefsteak tomatoes have a great number of small locules; and plum tomatoes have very few, very small locules.<ref>{{cite journal |last1=Muños |first1=Stéphane |last2=Ranc |first2=Nicolas |last3=Botton |first3=Emmanuel |last4=Bérard |first4=Aurélie |last5=Rolland |first5=Sophie |last6=Duffé |first6=Philippe |last7=Carretero |first7=Yolande |last8=Le Paslier |first8=Marie-Christine |last9=Delalande |first9=Corinne |last10=Bouzayen |first10=Mondher |last11=Brunel |first11=Dominique |last12=Causse |first12=Mathilde |display-authors=5 |title=Increase in Tomato Locule Number Is Controlled by Two Single-Nucleotide Polymorphisms Located Near ''WUSCHEL'' |journal=Plant Physiology |volume=156 |issue=4 |date=1 August 2011 |doi=10.1104/pp.111.173997 |doi-access=free |pages=2244–2254 |pmid=21673133 |pmc=3149950 |bibcode=2011PlanP.156.2244M }}</ref><ref>{{cite web |url=http://extensionpublications.unl.edu/assets/html/g1864/build/g1864.htm |title=Selecting Tomatoes for the Home Garden |publisher=University of Nebraska–Lincoln |access-date=20 November 2019 |archive-date=22 January 2020 |archive-url=https://web.archive.org/web/20200122140420/http://extensionpublications.unl.edu/assets/html/g1864/build/g1864.htm |url-status=live }}</ref><ref name="Lee-2007">{{cite journal |last1=Lee |first1=Eunkyung |last2=Sargent |first2=Steven A. |last3=Huber |first3=Donald J. |title=Physiological Changes in Roma-type Tomato Induced by Mechanical Stress at Several Ripeness Stages |journal=HortScience |volume=42 |issue=5 |year=2007 |pages=1237–1242 |doi=10.21273/HORTSCI.42.5.1237 |doi-access=free}}</ref> 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.<ref>{{cite web |url=https://www.gardenersworld.com/how-to/grow-plants/how-to-save-tomato-seed/ |author=<!--Staff writer(s); no by-line--> |title=How to save tomato seed|date=24 March 2019 |website=gardenersworld.com |publisher=Gardeners' World Magazine |access-date=3 January 2023 |url-status=live |archive-url=https://archive.today/20230103114534/https://www.gardenersworld.com/how-to/grow-plants/how-to-save-tomato-seed/ |archive-date=3 January 2023}}</ref>
The tomato has a mutualistic relationship with arbuscular mycorrhizal fungi such as ''Rhizophagus irregularis''. Scientists use the tomato as a model species for investigating such symbioses.<ref>{{cite journal |last1=Buendia |first1=Luis |last2=Wang |first2=Tongming |last3=Girardin |first3=Ariane |last4=Lefebvre |first4=Benoit |title=The LysM receptor-like kinase Sl LYK 10 regulates the arbuscular mycorrhizal symbiosis in tomato |journal=New Phytologist |date=April 2016 |volume=210 |issue=1 |pages=184–195 |doi=10.1111/nph.13753 |pmid=26612325 |doi-access=free|bibcode=2016NewPh.210..184B }}</ref>
<gallery class="center" mode="nolines" heights="180px" widths="180px"> File:Germinating tomatos.jpg |Seedlings 7 days after planting File:Tomato 27 days from planting seeds.jpg |27 days after planting File:Solanum lycopersicum - Flor tomaca 057.jpg|Flower File:Tomato fruit and flowers at day 52.jpg |52-day-old plant, first fruits File:Green Tomato.jpg|Unripe fruit on the vine </gallery>
=== Phylogeny === Like the potato, tomatoes belong to the genus ''Solanum'', which is a member of the nightshade family, the Solanaceae. That is a diverse family of flowering plants, often poisonous, that includes the mandrake (''Mandragora''), deadly nightshade (''Atropa''), and tobacco (''Nicotiana''), as shown in the outline phylogenetic tree (many branches omitted).<ref>Olmstead, Richard G., et al. "Phylogeny and provisional classification of the Solanaceae based on chloroplast DNA." Solanaceae IV 1.1 (1999): 1–137. https://www.researchgate.net/profile/Tharindu-Ranasinghe-2/post/Is-there-a-complete-phylogenetic-description-of-the-Solanaceae-family/attachment/59d63db579197b807799a764/AS%3A421051545735172%401477397919618/download/PHYLOGENY+AND+PROVISIONAL+CLASSIFICATION+OF+THE+SOLANACEAE+BASED+ON+CHLOROPLAST+DNA.pdf {{Webarchive|url=https://web.archive.org/web/20241009212631/https://www.researchgate.net/profile/Tharindu-Ranasinghe-2/post/Is-there-a-complete-phylogenetic-description-of-the-Solanaceae-family/attachment/59d63db579197b807799a764/AS%3A421051545735172%401477397919618/download/PHYLOGENY%20AND%20PROVISIONAL%20CLASSIFICATION%20OF%20THE%20SOLANACEAE%20BASED%20ON%20CHLOROPLAST%20DNA.pdf |date=9 October 2024 }}</ref>
{{clade |label1=Solanaceae |1={{clade |1= many garden flowers and other species |2={{clade |1=''Nicotiana'' (tobacco) |2={{clade |1=''Atropa'' (nightshades) |2={{clade |1=''Mandragora'' (mandrake) |2={{clade |label1=''Capsicum'' |1= (sweet and bell peppers) |label2=''Solanum'' |2={{clade |1='''''S. lycopersicum'' (tomato)''' |2=''S. tuberosum'' (potato) }} }} }} }} }} }} }}
=== Taxonomy ===
In 1753, Linnaeus placed the tomato in the genus ''Solanum'' (alongside the potato) as ''Solanum lycopersicum''. In 1768, Philip Miller moved it to its own genus, naming it ''Lycopersicon esculentum''.<ref>{{cite web |url=http://www.ipni.org/ipni/idPlantNameSearch.do?id=146898-2 |title=Lycopersicon esculentum |publisher=International Plant Name Index |archive-url=https://web.archive.org/web/20110606004439/http://www.ipni.org/ipni/idPlantNameSearch.do?id=146898-2 |archive-date=6 June 2011}}</ref> The name came into wide use, but was technically in breach of the plant naming rules because Linnaeus's species name ''lycopersicum'' still had priority. Although the name ''Lycopersicum lycopersicum'' was suggested by Karsten (1888), it is not used because it violates the International Code of Nomenclature<ref>{{cite web |url=http://www.iapt-taxon.org/nomen/main.php?page=art23 |publisher=International Association for Plant Taxonomy |title=International Code of Nomenclature for algae, fungi, and plants |access-date=14 September 2016 |archive-date=3 April 2021 |archive-url=https://web.archive.org/web/20210403051810/https://www.iapt-taxon.org/nomen/main.php?page=art23 |url-status=live }}</ref> barring the use of tautonyms in botanical nomenclature. The corrected name ''Lycopersicon lycopersicum'' (Nicolson 1974) was technically valid, because Miller's genus name and Linnaeus's species name differ in exact spelling. As ''Lycopersicon esculentum'' has become so well known, it was officially listed as a ''nomen conservandum'' in 1983, and would be the correct name for the tomato in classifications which do not place the tomato in the genus ''Solanum''.<ref>{{cite book |last=Turland |first=Nicholas |title=The Code Decoded: A user's guide to the International Code of Nomenclature for algae, fungi, and plants |url=https://plants.sdsu.edu/plantsystematics/pdfs/Turland2019-Nomenclature-ICN.pdf |publisher=Pensoft |date=2019 |page=85 |archive-date=9 October 2024 |access-date=9 October 2024 |archive-url=https://web.archive.org/web/20241009131253/https://plants.sdsu.edu/plantsystematics/pdfs/Turland2019-Nomenclature-ICN.pdf |url-status=live }}</ref>
Genetic evidence shows that Linnaeus was correct to put the tomato in the genus ''Solanum'', making ''S. lycopersicum'' the correct name.<ref name=NHM /><ref name="Peralta-2001">{{cite journal |first1=I.E. |last1=Peralta |first2=D.M. |last2=Spooner |title=Granule-bound starch synthase (GBSSI) gene phylogeny of wild tomatoes (''Solanum'' L. section ''Lycopersicon'' (Mill.) Wettst. subsection ''Lycopersicon'') |journal=American Journal of Botany |year=2001 |volume=88 |issue=10 |pages=1888–1902 |jstor=3558365 |pmid=21669622 |doi=10.2307/3558365 |doi-access=free|bibcode=2001AmJB...88.1888P }}</ref> 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, providing evidence of the close relationship between these species.<ref>{{Cite journal |author1=Jacobsen, E. |author2=Daniel, M.K. |author3=Bergervoet-van Deelen, J.E.M. |author4=Huigen, D.J. |author5=Ramanna, M.S. |s2cid=1015489 |year=1994 |title=The first and second backcross progeny of the intergeneric fusion hybrids of potato and tomato after crossing with potato |journal=Theoretical and Applied Genetics |volume=88 |issue=2 |pages=181–186 |doi=10.1007/BF00225895 |pmid=24185924}}</ref> Newer genomic studies have found that the tomato and the potato are very close relatives, forming a tight clade within ''Solanum''. A 2025 study suggests that the potato lineage may have been created by hybridization of a plant from the tomato lineage (not necessarily the modern tomato species) with a plant from the ''S. etuberosum'' lineage.<ref name=Zhang25>{{cite journal |last1=Zhang |first1=Zhiyang |last2=Zhang |first2=Pingxian |last3=Ding |first3=Yiyuan |last4=Wang |first4=Zefu |last5=Ma |first5=Zhaoxu |last6=Gagnon |first6=Edeline |last7=Jia |first7=Yuxin |last8=Cheng |first8=Lin |last9=Bao |first9=Zhigui |last10=Liu |first10=Zinan |last11=Wu |first11=Yaoyao |last12=Hu |first12=Yong |last13=Lian |first13=Qun |last14=Lin |first14=Weichao |last15=Wang |first15=Nan |last16=Ye |first16=Keyi |last17=Wang |first17=Hongru |last18=Zhang |first18=Jinzhe |last19=Zhou |first19=Yongfeng |last20=Liu |first20=Liang |last21=Li |first21=Suhua |last22=Lucas |first22=William J. |last23=Särkinen |first23=Tiina |last24=Knapp |first24=Sandra |last25=Rieseberg |first25=Loren H. |last26=Liu |first26=Jianquan |last27=Huang |first27=Sanwen |display-authors=5 |title=Ancient hybridization underlies tuberization and radiation of the potato lineage |journal=Cell |date=July 2025 |volume=188 |issue=19 |pages=5249–5265.e15 |doi=10.1016/j.cell.2025.06.034|pmid=40749684 |doi-access=free}}</ref>
== Plant breeding ==
=== Genetics ===
{{Infobox genome | image = <!-- Karyotype, for instance --> | caption = | taxId = GCF_000188115.5 | specimen = Heinz 1706 | ploidy = haploid | chromosomes = 12 | size = 827.4 Mb | genes = 31,217 | protein-coding = 25,557 | year = 2018 }}
An international consortium of researchers from 10 countries began sequencing the tomato genome in 2004.<ref>{{cite web |url=http://solgenomics.net/about/tomato_project_overview.pl |title=International Tomato Genome Sequencing Project |publisher=Sol Genomics Network |first=L. |last=Mueller |access-date=21 October 2009 |archive-date=24 July 2011 |archive-url=https://web.archive.org/web/20110724072309/http://solgenomics.net/about/tomato_project_overview.pl |url-status=live }}</ref><ref>{{Cite news |url=https://www.news.cornell.edu/stories/Jan07/SolanacaeNSF.kr.html |archive-url=https://web.archive.org/web/20100713035349/http://www.news.cornell.edu/stories/Jan07/SolanacaeNSF.kr.html |archive-date=13 July 2010 |title=Tomato genome project gets $1.8M |publisher=News.cornell.edu |last=Ramanujan |first=K. |date=30 January 2007 |access-date=27 October 2008}}</ref> A prerelease version of the genome was made available in December 2009.<ref>{{Cite web|url=http://solgenomics.net/tomato/|title=Tomato Genome Shotgun Sequence Prerelease}}</ref> The complete genome for the cultivar Heinz 1706 was published on 31 May 2012 in ''Nature''.<ref>{{Cite journal |last1=Sato |first1=S. |last2=Tabata |first2=S. |last3=Hirakawa |first3=H. |last4=Asamizu |first4=E. |last5=Shirasawa |first5=K. |last6=Isobe |first6=S. |last7=Kaneko |first7=T. |last8=Nakamura |first8=Y. |last9=Shibata |first9=D. |last10=Aoki |doi=10.1038/nature11119 |first10=K. |last11=Egholm |first11=M. |last12=Knight |first12=J. |last13=Bogden |first13=R. |last14=Li |first14=C. |last15=Shuang |first15=Y. |last16=Xu |first16=X. |last17=Pan |first17=S. |last18=Cheng |first18=S. |last19=Liu |first19=X. |last20=Ren |first20=Y. |last21=Wang |first21=J. |last22=Albiero |first22=A. |last23=Dal Pero |first23=F. |last24=Todesco |first24=S. |last25=Van Eck |first25=J. |last26=Buels |first26=R.M. |last27=Bombarely |first27=A. |last28=Gosselin |first28=J.R. |last29=Huang |first29=M. |last30=Leto |first30=J.A. |display-authors=5 |title=The tomato genome sequence provides insights into fleshy fruit evolution |journal=Nature |volume=485 |issue=7400 |pages=635–641 |year=2012 |pmid=22660326 |pmc=3378239 |bibcode=2012Natur.485..635T}}</ref><ref>{{cite news |url=http://www.rdmag.com/News/Feeds/2012/06/general-sciences-tomato-genome-is-sequenced-for-the-first-time/ |title=Tomato genome is sequenced for the first time |date=1 June 2012 |archive-url=https://web.archive.org/web/20120604005502/http://www.rdmag.com/News/Feeds/2012/06/general-sciences-tomato-genome-is-sequenced-for-the-first-time/ |archive-date=4 June 2012 |url-status=dead |work=R&D}}</ref> The latest reference genome published in 2021 had 799 MB and encodes 34,384 (predicted) proteins, spread over 12 chromosomes.<ref>{{Cite journal |display-authors=5 |last1=Su |first1=Xiao |last2=Wang |first2=Baoan |last3=Geng |first3=Xiaolin |last4=Du |first4=Yuefan |last5=Yang |first5=Qinqin |last6=Liang |first6=Bin |last7=Meng |first7=Ge |last8=Gao |first8=Qiang |last9=Yang |first9=Wencai |last10=Zhu |first10=Yingfang |last11=Lin |first11=Tao |date=15 December 2021 |title=A high-continuity and annotated tomato reference genome |journal=BMC Genomics |volume=22 |issue=1 |pages=898 |doi=10.1186/s12864-021-08212-x |issn=1471-2164 |pmc=8672587 |pmid=34911432 |doi-access=free}}</ref>
The first commercially available genetically modified food was a tomato called Flavr Savr, which was engineered to have a longer shelf life.<ref>{{ Cite book |last1=Redenbaugh |first1=K. |author2=Hiatt, B. |author3=Martineau, B. |author4=Kramer, M. |author5=Sheehy, R. |author6=Sanders, R. |author7=Houck, C. |author8=Emlay, D. |display-authors=5 |year=1992 |title=Safety Assessment of Genetically Engineered Fruits and Vegetables: A Case Study of the Flavr Savr Tomato |publisher=CRC Press |page=288 }}</ref> It could be vine ripened without compromising shelf life, which was expected to improve the flavor over ethylene-ripened tomatoes. However, it was not firmer than its unmodified parent, to the disappointment of its creators, who originally wanted to create a vine-ripened tomato that can survive machine-picking.<ref>Martineau, Belinda. 2001. ''First Fruit: The Creation of the Flavr Savr Tomato and the Birth of Biotech Food''. McGraw-Hill.</ref> The parent variety was also subpar in terms of yields.<ref>{{cite web |last1=Philippidis |first1=Alex |title=Mistakes Shorten Flavr Savr's Shelf Life |url=https://www.genengnews.com/insights/mistakes-shorten-first-approved-gmos-shelf-life/ |website=GEN - Genetic Engineering and Biotechnology News |date=12 April 2016 |access-date=4 August 2025 |archive-date=8 October 2025 |archive-url=https://web.archive.org/web/20251008042507/https://www.genengnews.com/insights/mistakes-shorten-first-approved-gmos-shelf-life/ |url-status=live }}</ref> As a result, the product was not commercially successful, and was sold only until 1997.<ref>{{cite journal |publisher=University of California, Agriculture and Natural Resources |journal=California Agriculture |volume=54 |issue=4 |pages=6–7 |doi=10.3733/ca.v054n04p6 |year=2000 |title=The case of the FLAVR SAVR tomato |last1=Bruening |first1=G. |last2=Lyons |first2=J.M. |doi-broken-date=10 July 2025 |doi-access=free}}</ref>
When the Mesoamericans domesticated tomatoes, they selected for less bitter fruits. This corresponded to the increased activity of a 2-oxoglutarate-dependent dioxygenase called ''23DOX'' (synonym ''GAME31'') during fruit ripening, which converts the bitter and slightly toxic α-tomatine into hydroxytomatine, which is eventually converted into the non-bitter and non-toxic esculeoside A.<ref name=Cardenas19>{{cite journal |last1=Cárdenas |first1=P.D. |last2=Sonawane |first2=P.D. |last3=Heinig |first3=U. |last4=Jozwiak |first4=A. |last5=Panda |first5=S. |last6=Abebie |first6=B. |last7=Kazachkova |first7=Y. |last8=Pliner |first8=M. |last9=Unger |first9=T. |last10=Wolf |first10=D |last11=Ofner |first11=I. |last12=Vilaprinyo |first12=E. |last13=Meir |first13=S. |last14=Davydov |first14=O. |last15=Gal-On |first15=A. |last16=Burdman |first16=S. |last17=Giri |first17=A. |last18=Zamir |first18=D. |last19=Scherf |first19=T. |last20=Szymanski |first20=J. |last21=Rogachev |first21=I. |last22=Aharoni |first22=A. |display-authors=5 |title=Pathways to defense metabolites and evading fruit bitterness in genus Solanum evolved through 2-oxoglutarate-dependent dioxygenases |journal=Nature Communications |date=14 November 2019 |volume=10 |issue=1 |pages=5169 |doi=10.1038/s41467-019-13211-4 |pmid=31727889|pmc=6856131 |bibcode=2019NatCo..10.5169C |doi-access=free}}</ref>
=== Commercial breeding ===
thumb|left|upright=1.5|Evolution of genetic diversity in cultivated tomatoes<ref>{{cite journal | last1=Schouten | first1=H. J. | last2=Tikunov | first2=Y. | last3=Verkerke | first3=W. | last4=Finkers | first4=R. | last5=Bovy | first5=A. | last6=Bai | first6=Y. | last7=Visser | first7=R. G. | title=Breeding Has Increased the Diversity of Cultivated Tomato in the Netherlands | journal=Frontiers in Plant Science | date=2019 | volume=10 | article-number=1606 | doi=10.3389/fpls.2019.01606 | doi-access=free | pmid=31921253 | pmc=6932954 | bibcode=2019FrPS...10.1606S }}</ref>
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 <!--lowercase-->"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 un-crossbred varieties. Before this, most tomatoes produced more sugar during ripening, and were sweeter and more flavorful.<ref name="Powell-2012">{{cite journal |last1=Powell |first1=Ann L.T. |last2=Nguyen |first2=Cuong V. |last3=Hill |first3=Theresa |last4=Cheng |first4=KaLai Lam |last5=Figueroa-Balderas |first5=Rosa |last6=Aktas |first6=Hakan |last7=Ashrafi |first7=Hamid |last8=Pons |first8=Clara |last9=Fernández-Muñoz |first9=Rafael |last10=Vicente |first10=Ariel |last11=Lopez-Baltazar |first11=Javier |last12=Barry |first12=Cornelius S. |last13=Liu |first13=Yongsheng |last14=Chetelat |first14=Roger |last15=Granell |first15=Antonio |last16=Van Deynze |first16=Allen |last17=Giovannoni |first17=James J. |last18=Bennett |first18=Alan B. |display-authors=5 |title=''Uniform ripening'' Encodes a ''Golden 2-like'' Transcription Factor Regulating Tomato Fruit Chloroplast Development |journal=Science |publisher=American Association for the Advancement of Science (AAAS) |volume=336 |issue=6089 |date=2012-06-29 |doi=10.1126/science.1222218 |pages=1711–1715 |pmid=22745430 |s2cid=23517955 |bibcode=2012Sci...336.1711P|hdl=10251/87703 |hdl-access=free }}</ref><ref name="Kolata-2012">{{cite news |last=Kolata |first=Gina |title=Flavor Is Price of Scarlet Hue of Tomatoes, Study Finds |url=https://www.nytimes.com/2012/06/29/science/flavor-is-the-price-of-tomatoes-scarlet-hue-geneticists-say.html |access-date=29 June 2012 |newspaper=The New York Times |date=28 June 2012 |archive-date=24 February 2021 |archive-url=https://web.archive.org/web/20210224031635/https://www.nytimes.com/2012/06/29/science/flavor-is-the-price-of-tomatoes-scarlet-hue-geneticists-say.html |url-status=live }}</ref>
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 mutation encodes a factor that produces defective chloroplasts with lower density in developing fruit, making them a lighter green, and reducing sugar in the resulting ripe fruit by 10–15%. Perhaps more importantly, the fruit chloroplasts are remodelled during ripening into chlorophyll-free chromoplasts that synthesize and accumulate the carotenoids lycopene, β-carotene, and other metabolites that are sensory and nutritional assets of the ripe fruit. The potent chloroplasts in the dark-green shoulders of the <!--uppercase-->"U" phenotype are beneficial here, but have the disadvantage of leaving green shoulders near the stems of the ripe fruit, and even cracked yellow shoulders. This is apparently because of oxidative stress due to overload of the photosynthetic chain in direct sunlight at high temperatures. Hence, genetic design of a commercial variety that combines the advantages of types "u" and "U" requires fine tuning, but may be feasible.<ref name="Cocaliadis-2014">{{cite journal |title=Increasing tomato fruit quality by enhancing fruit chloroplast function. A double-edged sword? |journal=Journal of Experimental Botany |date=10 April 2014 |volume=65 |issue=16 |pages=4589–4598 |doi=10.1093/jxb/eru165 |pmid=24723405 |doi-access=free |last1=Cocaliadis |first1=Maria Florencia |last2=Fernández-Muñoz |first2=Rafael |last3=Pons |first3=Clara |last4=Orzaez |first4=Diego |last5=Granell |first5=Antonio |hdl=10251/79375 |hdl-access=free}}</ref>
Breeders strive to produce tomato plants with improved yield, shelf life, size, and resistance to environmental pressures, including disease.<ref name="Klee-2018">{{cite journal |last1=Klee |first1=Harry J. |last2=Tieman |first2=Denise M. |s2cid=736072 |title=The genetics of fruit flavour preferences |journal=Nature |date=2018 |volume=19 |issue=6 |pages=347–356 |doi=10.1038/s41576-018-0002-5 |pmid=29563555}}</ref><ref name="Stevens-1986">{{cite book |last1=Stevens |first1=M. Allen |chapter=Inheritance of Tomato Fruit Quality Components |title=Plant Breeding Reviews |date=1986 |volume=4 |pages=273–311 |location=Westport, CT |publisher=Avi Publishing Company |doi=10.1002/9781118061015.ch9 |isbn=9781118061015}}</ref> These efforts have yielded unintended negative consequences on various fruit attributes. For instance, linkage drag, the introduction of an undesired trait during backcrossing, has altered the metabolism of the fruit. This trait is physically close to the desired allele along the chromosome. Breeding for traits like larger fruit has thus unintentionally altered nutritional value and flavor.<ref name="Klee-2018"/>
Breeders have turned to wild tomato species as a source of alleles to introduce beneficial traits into modern varieties. For example, wild relatives may possess higher amounts of fruit solids (associated with greater sugar content), or resistance to diseases such as to the early blight pathogen ''Alternaria solani''. However, this tactic has limitations, since selection for traits such as pathogen resistance can negatively impact other favorable traits such as fruit production.<ref name="Stevens-1986"/><ref>{{cite journal |last1=Chaerani |first1=Reni |last2=Voorrips |first2=Roeland E. |s2cid=36002406 |title=Tomato early blight (''Alternaria solani''): the pathogen, genetics, and breeding for resistance |journal=Journal of General Plant Pathology |date=2006 |volume=72 |issue=6 |pages=335–347 |doi=10.1007/s10327-006-0299-3 |bibcode=2006JGPP...72..335C}}</ref>
== Cultivation ==
{{Main list |List of tomato cultivars}}
The tomato is grown worldwide for its edible fruits, with thousands of cultivars.<ref>{{cite web |url=http://www.learnaboutag.org/resources/fact/tomatoes.pdf |title=Processing tomatoes |publisher=California Foundation for Agriculture in the Classroom |work=Commodity Fact Sheet |date=September 2014 |access-date=3 June 2016 |archive-date=11 June 2016 |archive-url=https://web.archive.org/web/20160611145322/http://www.learnaboutag.org/resources/fact/tomatoes.pdf |url-status=dead}}</ref>
<gallery class="center" mode="nolines" heights="180" widths="180"> File:Tomatoes in a market in France.jpg|Heirloom varieties in a French market File:Heirlooms.jpg|Heirloom cultivars Brandywine (biggest red), Black Krim (lower left) and Green Zebra (top left) File:Tomatoes for sale in France.jpg|Modern varieties in France File:Yellow cherry tomatoes.jpg|Yellow cherry tomatoes </gallery>
=== Hydroponic and greenhouse cultivation ===
Greenhouse tomato production in large-acreage commercial greenhouses and owner-operator stand-alone or multiple-bay greenhouses is increasing, providing fruit during those times of the year when field-grown fruit is not readily available. Smaller 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.<ref>{{cite web |last=Jones |first=J. Benton |title=Growing in the Greenhouse |url=http://www.growtomatoes.com/growing-in-the-greenhouse/ |work=Growing Tomatoes |archive-url=https://web.archive.org/web/20160520215357/http://www.growtomatoes.com/growing-in-the-greenhouse/ |access-date=14 August 2012 |archive-date=20 May 2016}}</ref> Tomatoes are also grown using hydroponics.<ref>{{Cite news |url=https://bio-protocol.org/bio101/e3121 |title=A simplified hydroponic culture of ''Arabidopsis'' |work=Bio-101 |access-date=4 Mar 2020 |archive-date=4 June 2020 |archive-url=https://web.archive.org/web/20200604154747/https://bio-protocol.org/bio101/e3121 |url-status=live }}</ref>
<gallery class="center" mode="nolines" heights="180" widths="180"> File:Tomato flower and young fruit.jpg|Flower and young fruit File:Tomato scanned.jpg|Flowers and ripe fruit can be present simultaneously. File:Tomato P5260299b.jpg|Hydroponic cultivation File:Greenhouses near El Ejido.jpg|Greenhouse cultivation in Andalusia </gallery>
=== Picking and ripening ===
To facilitate transportation and storage, tomatoes are often picked unripe (green) and ripened in storage with the plant hormone ethylene.<ref>{{cite web |last1=Russell |first1=Adam |title=Pick tomatoes at color break: Ripening off the vine extends harvest, quality with no taste difference |url=https://agrilifetoday.tamu.edu/2022/06/10/pick-tomatoes-at-color-break/ |website=Agrilife Today |access-date=22 October 2024 |date=10 June 2022 |quote=Tomatoes ripen as they begin to produce ethylene gas, which promotes the process. ... many commercially grown tomatoes are picked green for shipping, then treated with ethylene gas or placed in "ripening rooms" to promote ripening.}}</ref> At industrial scale, such as for canning, tomatoes are picked mechanically. The machine cuts the whole vine and uses sensors to separate ripe tomatoes from the rest of the plant, which is returned to the farm for use either as green manure or to be grazed by livestock.<ref name="Bittman-2013">{{cite news |title=Not All Industrial Food Is Evil |url=http://opinionator.blogs.nytimes.com/2013/08/17/not-all-industrial-food-is-evil/ |access-date=18 August 2013 |newspaper=The New York Times |date=17 August 2013 |last=Bittman |first=Mark |archive-date=29 August 2013 |archive-url=https://web.archive.org/web/20130829184407/http://opinionator.blogs.nytimes.com/2013/08/17/not-all-industrial-food-is-evil/ |url-status=live }}</ref>
=== Production === {{owidslider |start = 2023 |list = Template:OWID/tomato production#gallery |location = commons |caption = |title = |language = |file = link=|thumb|upright=1.6|Tomato production |startingView = World }}
{{Table alignment}} {| class="wikitable floatright col2right" |+ Tomato production <br>{{small|2023, millions of tonnes}}<br/> |- |{{CHN}} ||70.1 |- |{{IND}} ||29.4 |- |{{TUR}} ||13.3 |- |{{USA}} ||12.4 |- |{{EGY}} ||6.2 |- |{{MEX}} ||4.4 |- |'''World''' ||'''192.3''' |- |colspan=2 |{{small|Source: FAOSTAT<<br> of the United Nations}}<ref name="faostat">{{cite web |url=https://www.fao.org/faostat/en/#data/QCL |title=Tomato production in 2023, Crops/Regions/World list/Production Quantity/Year (pick lists) |date=2025 |publisher=UN Food and Agriculture Organization, Corporate Statistical Database (FAOSTAT) |access-date=12 August 2025 |archive-date=12 November 2016 |archive-url=https://web.archive.org/web/20161112130804/https://www.fao.org/faostat/en/#data/QCL |url-status=live }}</ref> |}
In 2023, world production of tomatoes was 192 million tonnes, led by China with 36% of the total, followed by India, Turkey, and the United States as secondary producers (table).
== Pests and diseases ==
=== Pests ===
Common tomato pests include the tomato bug, stink bugs, cutworms, tomato hornworms and tobacco hornworms, aphids, cabbage loopers, whiteflies, tomato fruitworms, flea beetles, red spider mite, ''Tuta absoluta'' (tomato leafminer), slugs,<ref>{{Cite web |url=http://www.extension.umn.edu/distribution/horticulture/dg7561.html |title=Slugs in Home Gardens |last1=Hahn, J. |last2=Fetzer, J. |year=2009 |publisher=University of Minnesota Extension |archive-url=https://web.archive.org/web/20110311110055/http://www.extension.umn.edu/distribution/horticulture/dg7561.html |archive-date=11 March 2011 |access-date=23 June 2012}}</ref> and Colorado potato beetles. The tomato russet mite, ''Aculops lycopersici'', feeds on foliage and young fruit of tomato plants, causing shrivelling and necrosis of leaves, flowers, and fruit, possibly killing the plant.<ref name="ISC-2015">{{Cite web |url=http://www.cabi.org/isc/datasheet/56111 |title=Aculops lycopersici (tomato russet mite) |publisher=Invasive Species Compendium, Centre for Agriculture and Biosciences International |location=Wallingford, UK |date=23 June 2015 |access-date=11 November 2016 |archive-date=12 November 2016 |archive-url=https://web.archive.org/web/20161112020537/http://www.cabi.org/isc/datasheet/56111 |url-status=live }}</ref>
After an insect attack tomato plants produce systemin, a plant peptide hormone. This activates defensive mechanisms, such as the production of protease inhibitors to slow the growth of insects. The hormone was first identified in tomatoes.<ref name="Narvaez-Vasquez-2008">{{ cite book |last1=Narvaez-Vasquez |first1=J. |last2=Orozco-Cardenas |first2=M.L. |editor=Schaller, A. |chapter=15 Systemins and AtPeps: Defense-related Peptide Signals |year=2008 |title=Induced Plant Resistance to Herbivory |isbn=978-1-4020-8181-1 }}</ref>
<gallery class="center" mode="nolines" heights="115" widths="230"> File:Engytatus modestus closeup 2.jpg|Tomato bug feeding on plant sap File:Tomato fruitworm.jpg|Tomato fruitworm feeding on unripe fruit File:Tomato hornworm.jpg|Tomato hornworm larva on stem File:20230811 Aculops lycopersici 08 D.jpg|Tomato russet mites on greenhouse plant </gallery>
=== Diseases ===
{{Main list|List of tomato diseases}}
Tomato cultivars vary widely in their resistance to disease. Modern hybrids focus on improving disease resistance over the heirloom plants. A common tomato disease is tobacco mosaic virus. Handling cigarettes and other infected tobacco products can transmit the virus to tomato plants.<ref>{{cite web |url=http://www.extension.umn.edu/distribution/horticulture/DG1168.html |author1=Pfleger, F.L. |author2=Zeyen, R.J. |title=Tomato-Tobacco Mosaic Virus Disease |year=2008 |publisher=University of Minnesota Extension |access-date=23 June 2012 |url-status=dead |archive-url=https://web.archive.org/web/20120614075040/http://www.extension.umn.edu/distribution/horticulture/dg1168.html |archive-date=14 June 2012}}</ref> A serious 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.<ref>{{cite web |last=Goldberg |first=N.P. |url=https://pubs.nmsu.edu/_h/H106/index.html |title=Curly Top Virus: Guide H-106 |publisher=College of Agriculture, Consumer and Environmental Sciences. New Mexico State University |access-date=9 October 2024 |archive-date=9 October 2024 |archive-url=https://web.archive.org/web/20241009085745/https://pubs.nmsu.edu/_h/H106/index.html |url-status=live }}</ref> Bacterial wilt is another common disease impacting yield.<ref name="Fitzpatrick-2020">{{cite journal |last1=Fitzpatrick |first1=Connor R. |last2=Salas-González |first2=Isai |last3=Conway |first3=Jonathan M. |last4=Finkel |first4=Omri M. |last5=Gilbert |first5=Sarah |last6=Russ |first6=Dor |last7=Teixeira |first7=Paulo José Pereira Lima |last8=Dangl |first8=Jeffery L. |title=The Plant Microbiome: From Ecology to Reductionism and Beyond |journal=Annual Review of Microbiology |publisher=Annual Reviews |volume=74 |issue=1 |date=8 September 2020 |issn=0066-4227 |doi=10.1146/annurev-micro-022620-014327 |pages=81–100 |s2cid=219621296 |pmid=32530732 |url=https://cdr.lib.unc.edu/downloads/br86bc78d |doi-access=free |osti=1802632 |archive-date=14 August 2022 |access-date=29 July 2022 |archive-url=https://web.archive.org/web/20220814153916/https://cdr.lib.unc.edu/downloads/br86bc78d |url-status=live }}</ref> Wang ''et al.'', 2019 found phage combination therapies to reduce the impact of bacterial wilt, sometimes by reducing bacterial abundance and sometimes by selecting for resistant but slow growing genetics.<ref name="Fitzpatrick-2020"/>
<gallery class="center" mode="nolines" heights="200" widths="200"> File:Tomato with Phytophthora infestans (late blight).jpg|Late blight, caused by the oomycete ''Phytophthora infestans'' File:Ralstonis wilt symptom.jpg|Wilt caused by the bacterium<br />''Ralstonia solanacearum'' File:Tomaquera amb Fusarium HV.JPG|Wilt caused by ''Fusarium oxysporum'' File:Tomato (Solanum lycopersicum)- Root knot nematodes - 27421750599.jpg|Nematode root-knot caused by ''Meloidogyne incognita'' </gallery>
== As food ==
=== Culinary ===
{{further|List of tomato dishes}}
Tomatoes, with their umami flavor, are extensively used in Mediterranean cuisine as a key ingredient in pizza and many pasta sauces.<ref name="Fleming-2013">{{cite news |url=https://www.theguardian.com/lifeandstyle/wordofmouth/2013/apr/09/umami-fifth-taste |title=Umami: why the fifth taste is so important |last=Fleming |first=Amy |newspaper=The Guardian |date=9 April 2013 |access-date=18 February 2017 |archive-date=15 December 2023 |archive-url=https://web.archive.org/web/20231215214931/https://www.theguardian.com/lifeandstyle/wordofmouth/2013/apr/09/umami-fifth-taste |url-status=live }}</ref> Tomatoes are used in Spanish gazpacho<ref>{{cite web |website=Royal Spanish Academy |title=gazpacho |url=https://dle.rae.es/gazpacho?m=form |access-date=9 October 2024 |archive-date=14 August 2024 |archive-url=https://web.archive.org/web/20240814001023/https://dle.rae.es/gazpacho?m=form |url-status=live }}</ref> and Catalan {{lang|ca|pa amb tomàquet}}.<ref>{{cite news |title=Pa Amb Tomàquet (Catalan Tomato Bread) |url=http://www.foodandwine.com/recipes/catalan-tomato-bread |access-date=9 October 2024 |work=Food & Wine |archive-date=30 March 2023 |archive-url=https://web.archive.org/web/20230330175813/https://www.foodandwine.com/recipes/catalan-tomato-bread |url-status=live }}</ref> 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.<ref>{{Cite web |last=Beyer |first=Greg |date=7 Apr 2024 |title=The History of the Tomato: The Fruit that Spread Round the World |url=https://www.thecollector.com/history-of-tomato/ |access-date=10 June 2024 |website=The Collector}}</ref>
Tomatoes were gradually incorporated into Indian curry dishes after Europeans introduced them.{{sfn|Collingham|2006|p=165}} A Kashmiri curry, rogan josh, often contains tomato; it may originally have been colored red with chili pepper,<ref name="Singh-1973">{{cite book |last=Singh |first=Dharamjit |title=Indian Cookery |url=https://archive.org/details/indiancookery00sing/page/21 |year=1973 |publisher=Penguin |isbn=978-0140461411 |page=[https://archive.org/details/indiancookery00sing/page/21 21,58]}}</ref> and tomatoes may characterize the Punjabi version of the dish.<ref name="Bhangal-2013">{{cite book |last=Bhangal |first=Jasprit |title=Indian Cooking with Four Ingredients |year=2013 |publisher=Troubador |isbn=9781780884868 |page=101}}</ref> The modern British curry tikka masala often has a tomato and cream sauce.{{sfn|Collingham|2006|pp=1–11}}
<gallery class="center" mode="nolines" heights="180" widths="180"> File:Tomato soup with bread.jpg|Tomato soup with croutons File:Rogan josh02.jpg|Rogan josh, a curry often made with tomatoes Bloody Mary Coctail with celery stalk - Evan Swigart.jpg|Bloody Mary, a tomato cocktail File:Pan tumaca cortado (cropped).jpg|{{lang|ca|Pa amb tomàquet}}, Catalan tomato bread File:Tomates farcies végétariennes.jpg|Tomatoes stuffed with egg and Parmesan cheese </gallery>
=== Storage ===
Tomatoes keep best unwashed at room temperature and out of direct sunlight, rather than in a refrigerator.<ref>{{cite web |url=http://anrcatalog.ucdavis.edu/pdf/8116.pdf |title=Tomatoes:Safe Methods to Store, Preserve, and Enjoy |date=March 2004 |last1=Parnell |first1=Tracy L. |last2=Suslow |first2=Trevor V. |last3=Harris |first3=Linda J. |publisher=University of California: Division of Agriculture and Natural Resources |work=ANR Catalog |access-date=18 February 2013 |archive-date=14 February 2013 |archive-url=https://web.archive.org/web/20130214062847/http://anrcatalog.ucdavis.edu/pdf/8116.pdf |url-status=dead}}</ref><ref>{{cite web |url=http://ohioline.osu.edu/hyg-fact/5000/pdf/5532.pdf |title=Selecting, Storing and Serving Ohio Tomatoes, HYG-5532-93 |publisher=Ohio State University |access-date=27 October 2008}}</ref> Storing stem down can prolong shelf life.<ref>[https://web.archive.org/web/20130406050733/http://www.cooksillustrated.com/howto/print/detail.asp?docid=1173 How To Cook]. Cooks Illustrated (1 July 2008). Retrieved on 5 September 2013.</ref> Unripe tomatoes can be kept in a paper bag to ripen.<ref>{{cite web |url=http://www.cpma.ca/en/about/areas-of-focus/vegetables#Tomato |title=Vegetables |publisher=Canadian Produce Marketing Association |work=Canadian Produce Marketing Association Website |access-date=18 February 2013 |url-status=dead |archive-url=https://web.archive.org/web/20130405083130/http://www.cpma.ca/en/about/areas-of-focus/vegetables |archive-date=5 April 2013}}</ref> Tomatoes can be preserved by canning, freezing, drying, or cooking down to a paste or puree.<ref>{{cite web |last=Watson |first=Molly |title=How To Preserve Tomatoes: Canning, Drying, and Freezing Tomatoes With Ease |url=https://www.thespruceeats.com/how-to-preserve-tomatoes-2217665 |website=The Spruce Eats |date=26 July 2024 |access-date=9 October 2024}}</ref>
=== Nutrition ===
{{nutritionalvalue |name=Red tomatoes, raw |water=94.5 g |kJ=74 |protein=0.9 g |fat=0.2 g |carbs=3.9 g |fiber=1.2 g |sugars=2.6 g |calcium_mg=10 |iron_mg=0.27 |magnesium_mg=11 |phosphorus_mg=24 |potassium_mg=237 |manganese_mg=0.114 |sodium_mg=5 |zinc_mg=0.17 |vitC_mg=14 |thiamin_mg=0.037 |riboflavin_mg=0.019 |niacin_mg=0.594 |pantothenic_mg=0.089 |vitB6_mg=0.08 |folate_ug=15 |vitA_ug=42 |betacarotene_ug=449 |lutein_ug=123 |opt1n=Lycopene |opt1v=2570 μg |vitE_mg=0.54 |vitK_ug=7.9 |note=[https://fdc.nal.usda.gov/food-details/170457/nutrients Link to USDA Database entry] }}
A raw tomato is 95% water, 4% carbohydrates, and less than 1% each of fat and protein (table). In a reference amount of {{convert|100|g|oz}}, raw tomatoes supply 18 calories and 16% of the Daily Value of vitamin C, but otherwise have low micronutrient content (table).
=== Effects on health ===
The US Food and Drug Administration has determined there is little credible evidence that tomatoes or tomato-based foods reduce the risk of various types of cancer.<ref name="Kavanaugh-2007">{{cite journal |last1=Kavanaugh |first1=Claudine J. |last2=Trumbo |first2=Paula R. |last3=Ellwood |first3=Kathleen C. |title=The U.S. Food and Drug Administration's evidence-based review for qualified health claims: tomatoes, lycopene, and cancer |journal=Journal of the National Cancer Institute |volume=99 |issue=14 |pages=1074–85 |date=July 2007 |pmid=17623802 |doi=10.1093/jnci/djm037 |url=https://academic.oup.com/jnci/article/99/14/1074/937795 |url-access=subscription |archive-date=26 November 2022 |access-date=26 October 2024 |archive-url=https://web.archive.org/web/20221126041616/https://academic.oup.com/jnci/article/99/14/1074/937795 |url-status=live }}</ref>
In a 2011 scientific review, the European Food Safety Authority concluded that lycopene did not favorably influence DNA, skin exposed to ultraviolet radiation, heart function or vision.<ref name="EFSA-2011">{{cite journal |title=Scientific Opinion on the substantiation of health claims related to lycopene and protection of DNA, proteins and lipids from oxidative damage (ID 1608, 1609, 1611, 1662, 1663, 1664, 1899, 1942, 2081, 2082, 2142, 2374), protection of the skin from UV-induced (including photo-oxidative) damage (ID 1259, 1607, 1665, 2143, 2262, 2373), contribution to normal cardiac function (ID 1610, 2372), and maintenance of normal vision (ID 1827) pursuant to Article 13(1) of Regulation (EC) No 1924/2006 |author=European Food Safety Authority |journal=EFSA Journal |year=2011 |volume=9 |issue=4 |pages=2031 |doi=10.2903/j.efsa.2011.2031 |doi-access=free}}</ref>
=== Toxins ===
The leaves, stem, and green unripe fruit of the tomato plant contain small amounts of the alkaloid tomatine.<ref name="Mcgee-2009"/> They contain small amounts of solanine, a toxic alkaloid found in larger amounts in potato leaves and other members of the nightshade family.<ref name="Barceloux-2009">{{Cite journal |last=Barceloux |first=D.G. |title=Potatoes, Tomatoes, and Solanine Toxicity (''Solanum tuberosum'' L., ''Solanum lycopersicum'' L.) |journal=Disease-a-Month |year=2009 |volume=55 |issue=6 |pages=391–402 |pmid=19446683 |doi=10.1016/j.disamonth.2009.03.009 |s2cid=41740029}}</ref><ref name=NIH>{{cite web |title=Executive Summary Chaconine and Solanine: 6.0 through 8.0 |url=https://ntp.niehs.nih.gov/sites/default/files/ntp/htdocs/chem_background/exsumpdf/chaconinesolanine_508.pdf |publisher=NIH |url-status=live |archive-url=https://web.archive.org/web/20140928132105/http://ntp.niehs.nih.gov/ntp/htdocs/chem_background/exsumpdf/chaconinesolanine_508.pdf |archive-date=28 September 2014}}</ref> Tomato plants can be toxic to dogs if they eat large amounts of the fruit, or chew plant material.<ref>{{cite book |title=Hound Health Handbook: The Definitive Guide to Keeping your Dog Happy |last=Brevitz |first=B. |page=404 |publisher=Workman Publishing Company |year=2004 |isbn=978-0-7611-2795-6 |url=https://archive.org/details/houndhealthhandb0000brev/page/404}}</ref>
Small amounts of tomato foliage are sometimes used for flavoring, and the green fruit of unripe red tomato varieties is sometimes used for cooking, particularly as fried green tomatoes.<ref name="Mcgee-2009">{{Cite news |last=Mcgee |first=H. |title=Accused, Yes, but Probably Not a Killer |newspaper=The New York Times |date=29 July 2009 |url=https://www.nytimes.com/2009/07/29/dining/29curi.html |access-date=26 March 2010 |archive-date=17 October 2016 |archive-url=https://web.archive.org/web/20161017194831/http://www.nytimes.com/2009/07/29/dining/29curi.html |url-status=live }}</ref>
=== ''Salmonella'' outbreaks ===
Tomatoes have been linked to multiple ''Salmonella'' food poisoning outbreaks in the US.<ref>{{cite web |url=http://www.foodsafety.ksu.edu/en/article-details.php?a=3&c=32&sc=419&id=953 |archive-url=https://web.archive.org/web/20080126104725/http://www.foodsafety.ksu.edu/en/article-details.php?a=3&c=32&sc=419&id=953 |url-status=dead |archive-date=26 January 2008 |title=A selection of North American tomato related outbreaks from 1990–2005 |publisher=Food Safety Network |date=30 October 2006 |access-date=20 July 2010}}</ref><ref>{{cite web |url=http://www.cbsnews.com/stories/2006/10/30/national/main2138331.shtml |archive-url=https://web.archive.org/web/20071013133735/http://www.cbsnews.com/stories/2006/10/30/national/main2138331.shtml |url-status=dead |archive-date=13 October 2007 |title=CDC Probes Salmonella Outbreak, Health Officials Say Bacteria May Have Spread Through Some Form Of Produce |work=CBS News |date=30 October 2006 |access-date=27 October 2008}}</ref> One in 2008 caused the temporary removal of tomatoes from stores and restaurants across the United States and parts of Canada.<ref>{{cite web |url=http://www.canada.com/calgaryherald/news/city/story.html?id=0c30bbc6-5fa0-41c2-9148-f57e622c0cdd |title=Tomatoes taken off menus |publisher=Calgary Herald |date=11 June 2008 |access-date=20 July 2010 |url-status=dead |archive-url=https://web.archive.org/web/20110825075739/http://www.canada.com/calgaryherald/news/city/story.html?id=0c30bbc6-5fa0-41c2-9148-f57e622c0cdd |archive-date=25 August 2011 |df=dmy-all}}</ref> In 2022 and 2023, an outbreak of Salmonella Senftenberg ST14 affected the US and 12 countries in Europe.<ref name="ECDPC-2023">{{cite web |title=Multi-country outbreak of Salmonella Senftenberg ST14 infections possibly linked to cherry-like tomatoes |url=https://www.ecdc.europa.eu/en/publications-data/multi-country-outbreak-salmonella-senftenberg-st14-infections |publisher=European Centre for Disease Prevention and Control |access-date=22 October 2024 |date=27 July 2023 |archive-date=8 December 2024 |archive-url=https://web.archive.org/web/20241208095711/https://www.ecdc.europa.eu/en/publications-data/multi-country-outbreak-salmonella-senftenberg-st14-infections |url-status=live }}</ref>
== In popular culture ==
=== Celebrations ===
[[File:Tomatotree.JPG|thumb|The "tomato tree" at the Walt Disney World Resort's experimental greenhouses<ref name="Guinness-2006"/>]]
A massive "tomato tree" in the Walt Disney World Resort's experimental greenhouses in Lake Buena Vista, Florida may have been the largest single tomato plant. It yielded a harvest of more than 32,000 tomatoes, together weighing {{cvt|522|kg|lb}}.<ref name="Guinness-2006">{{cite web |url=https://www.guinnessworldrecords.com/world-records/most-tomatoes-harvested-from-one-plant-in-one-year |title=Most tomatoes harvested from one plant in one year |website=Guinness World Records |date=20 April 2006 |access-date=22 June 2023 |archive-url=https://web.archive.org/web/20221205152127/https://www.guinnessworldrecords.com/world-records/most-tomatoes-harvested-from-one-plant-in-one-year |archive-date=5 December 2022 |url-status=live}}</ref><ref name="Disney">[https://web.archive.org/web/20070717063219/http://wdwnews.com/ViewImage.aspx?ImageID=101932 The country's only single vine "tomato tree" growing in The Land pavilion at Epcot]. Walt Disney World News</ref>
The town of Buñol, Spain, annually celebrates La Tomatina, a festival centered on an enormous tomato fight. On 30 August 2007, as many as 40,000 Spaniards gathered to throw {{cvt |115000 |kg |lb}} of tomatoes at each other in the festival.<ref>{{cite web |url=http://itn.co.uk/news/9a5a1671ceba4f43741dc008f237c1ea.html |archive-url=https://web.archive.org/web/20071012035247/http://itn.co.uk/news/9a5a1671ceba4f43741dc008f237c1ea.html |archive-date=12 October 2007 |title=Spain's tomato fighters see red |publisher=ITV |date=30 August 2007 |access-date=2 April 2009}}</ref>
Some US states have adopted the tomato as a state fruit or vegetable. 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.<ref>{{cite web |title=Vine Ripe Pink Tomato |url=https://statesymbolsusa.org/symbol-official-item/arkansas/state-food-agriculture-symbol/south-arkansas-vine-ripe-pink-tomato |website=State Symbols USA |date=12 May 2014 |access-date=9 October 2024 |archive-date=9 October 2024 |archive-url=https://web.archive.org/web/20241009140710/https://statesymbolsusa.org/symbol-official-item/arkansas/state-food-agriculture-symbol/south-arkansas-vine-ripe-pink-tomato |url-status=live }}</ref> In 2009, the state of Ohio passed a law making the tomato the state's official fruit, while tomato juice has been the state's official beverage since 1965.<ref>{{cite web |title=Tomato |url=https://statesymbolsusa.org/symbol-official-item/ohio/state-food-agriculture-symbol/tomato |website=State Symbols USA |date=5 September 2014 |access-date=9 October 2024 |archive-date=9 October 2024 |archive-url=https://web.archive.org/web/20241009140711/https://statesymbolsusa.org/symbol-official-item/ohio/state-food-agriculture-symbol/tomato |url-status=live }}</ref> Livingston's plant breeding is commemorated in his home town of Reynoldsburg with an annual Tomato Festival;<ref>{{cite web |title=Tomato Festival, Reynoldsburg, Ohio |url=https://www.reytomatofest.com/ |website=Reynoldsburg Tomato Festival |access-date=9 October 2024}}</ref> it calls itself "The Birthplace of the Tomato".<ref name="Reynoldsburg">[https://web.archive.org/web/20110111123746/http://www.ci.reynoldsburg.oh.us/about-reynoldsburg.aspx About Reynoldsburg]. ci.reynoldsburg.oh.us</ref> In Finland, the ''Tomatkarnevalen'' is held annually in the town of Närpes.<ref>{{cite web |title=Tomatkernevalen |url=https://tomatkarnevalen.fi/ |website=Tomatkernevalen |access-date=9 October 2024 |archive-date=9 October 2024 |archive-url=https://web.archive.org/web/20241009163104/https://tomatkarnevalen.fi/ |url-status=live }}</ref>
Tomatoes are sometimes thrown in public protests. Embracing it for this connotation, the Dutch Socialist party adopted the tomato as their logo.<ref>{{cite web |last1=Holligan |first1=Anna |title=Dutch election: Emile Roemer seduces with Socialist charms |url=https://www.bbc.co.uk/news/19408907 |publisher=BBC |access-date=22 October 2024 |date=30 August 2012 |quote=It still alludes to past Maoist allegiances with its emblem of a white star in a red tomato, harking back to the days when activists hurled fruit at opponents during protests.}}</ref> The same meaning is evoked in the name of the American review-aggregation website for film and television, "Rotten Tomatoes", though its founder mentions a scene in the 1992 movie ''Leolo'' as the immediate source of the name.<ref>{{cite magazine |url=https://www.wired.com/story/behind-the-scenes-rotten-tomatoes/ |url-status=dead |archive-url=https://web.archive.org/web/20200121151651/https://www.wired.com/story/behind-the-scenes-rotten-tomatoes/ |title=Behind the Scenes at Rotten Tomatoes |first=Simon |last=Van Zuylen-Wood |magazine=Wired |date=21 January 2020 |archive-date=21 January 2020}}</ref>
<gallery class="center" mode="nolines" heights="180" widths="180"> File:Arrojando tomates desde un camión - La Tomatina 2010.jpg|Throwing tomatoes from a truck during the Spanish ''Tomatina'' festival File:Tomatkarnevalen i Närpes 1993c.jpg|{{lang|fi|Tomatkarnevalen}} (The Tomato Carnival) in Närpes, Finland, in 1993 </gallery>
=== Fruit or vegetable ===
{{See also|Vegetable#Terminology}}
Although the tomato is cooked and eaten as a vegetable, botanically, a tomato is a fruit, specifically a berry, consisting of the ovary, together with its seeds, of a flowering plant.<ref>{{Cite book |last1=Michaels |first1=Tom |url=https://open.lib.umn.edu/horticulture/ |title=The Science of Plants |last2=Clark |first2=Matt |last3=Hoover |first3=Emily |last4=Irish |first4=Laura |last5=Smith |first5=Alan |last6=Tepe |first6=Emily |publisher=University of Minnesota Libraries Publishing |editor-last=Tepe |editor-first=Emily |language=en |chapter=Chapter 8.1 Fruit Morphology |date=20 June 2022 |isbn=9781946135872 |chapter-url=https://open.lib.umn.edu/horticulture/chapter/8-1-fruit-morphology/ |archive-date=25 April 2022 |access-date=25 April 2022 |archive-url=https://web.archive.org/web/20220425160853/https://open.lib.umn.edu/horticulture/ |url-status=live }}</ref><ref name="Abadi-2018">{{cite web |first=Mark |last=Abadi |title=A tomato is actually a fruit — but it's a vegetable at the same time |url=https://www.businessinsider.com/tomato-fruit-or-vegetable-2018-5 |work=Business Insider |access-date=21 November 2019 |date=26 May 2018 |archive-date=15 February 2020 |archive-url=https://web.archive.org/web/20200215004602/https://www.businessinsider.com/tomato-fruit-or-vegetable-2018-5 |url-status=live }}</ref> The issue has led to legal dispute in the United States. In 1887, U.S. tariff laws that imposed a duty on vegetables, but not on fruit, caused the tomato's status to become a matter of legal importance. In ''Nix v. Hedden'', the U.S. Supreme Court settled the controversy on 10 May 1893, by declaring that for the purposes of the Tariff of 1883 only, the tomato is a vegetable, based on the popular definition that classifies vegetables by use—they are generally served with dinner and not dessert.<ref>{{ussc |name=Nix v. Hedden |volume=149 |page=304 |year=1893}}.</ref>
== See also ==
{{div col |colwidth=22em}} * La Tomatina, world's largest tomato food fight * List of countries by tomato production * List of tomato dishes * Marglobe, an early attempt at breeding a disease-resistant tomato * Ring culture * Physalis, a similar fruit also used in cooking * Tomato effect * Tomato jam * Nightshades ** Potato ** Eggplant ** Tomatillo, a similar fruit from the related genus {{div col end}}
== References ==
{{Reflist |30em}}
== Sources == * {{cite book |last=Collingham |first=Lizzie |author-link=Lizzie Collingham |title=Curry: A Tale of Cooks and Conquerors |date=2006 |orig-year=2005 (Chatto & Windus) |publisher=Vintage Books |location=London |isbn=978-0-099-43786-4}} * {{cite book |last=Smith |first=A.F. |year=1994 |title=The Tomato in America: Early History, Culture, and Cookery |publisher=University of South Carolina Press |location=Columbia, South Carolina |isbn=978-1-57003-000-0 |url=https://archive.org/details/tomatoinamericae00smit_0}}
== Further reading ==
* Tracy, Will W. '' Tomato culture: A practical treatise on the tomato: '' (1907) [https://dn721908.ca.archive.org/0/items/tomatocultureapr28011gut/28011.txt online] primary source on expert knowledge in early 20th century
* David Gentilcore. ''Pomodoro! A History of the Tomato in Italy'' (Columbia University Press, 2010), scholarly history * {{cite journal |title=The Chemical Interactions Underlying Tomato Flavor Preferences |doi=10.1016/j.cub.2012.04.016 |journal=Current Biology |date=5 June 2012 |pages=1035–1039 |volume=22 |issue=11 |pmid=22633806 |last1=Tieman |first1=D. |last2=Bliss |first2=P. |last3=McIntyre |first3=L.M. |last4=Blandon-Ubeda |first4=A. |last5=Bies |first5=D. |last6=Odabasi |first6=A.Z. |last7=Rodríguez |first7=G.R. |last8=van der Knaap |first8=E |last9=Taylor |first9=M.G. |last10=Goulet |first10=C. |last11=Mageroy |first11=M.H. |last12=Snyder |first12=D.J. |last13=Colquhoun |first13=T. |last14=Moskowitz |first14=H. |last15=Clark |first15=D.G. |last16=Sims |first16=C. |last17=Bartoshuk |first17=L. |last18=Klee |first18=H.J. |doi-access=free |bibcode=2012CBio...22.1035T |ref=none}}
== External links ==
{{Commons category multi |Solanum lycopersicum |Tomatoes}} * {{Cookbook-inline |Tomato}} * {{Wikibooks inline |Horticulture}} * {{Wikibooks inline |Tomato}} * {{Wikispecies-inline |Solanum lycopersicum}} * {{Wiktionary-inline |tomato}} * [https://www.sgn.cornell.edu/about/tomato_sequencing.pl Tomato Genome Sequencing Project] – Sequencing of the twelve tomato chromosomes. * [http://www.eu-sol.wur.nl/ Tomato core collection database] – Phenotypes and images of 7,000 tomato cultivars
{{Tomatoes}}
{{Taxonbar |from1=Q23501 |from2=Q15530945}} {{Authority control}}
Tomatoes Category:Crops originating from Mexico Category:Crops originating from indigenous Americans Category:Crops originating from South America Category:Fruit vegetables Category:Fruits originating in North America Category:Fruits originating in South America Category:Plants described in 1753 Category:Solanum Category:Plants with compound leaves