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This article is about the fruit. For other uses, see Citrus (disambiguation).

Citrus
Temporal range: Tortonian–Present, 8–0 Ma[1]
Sweet orange (Citrus × sinensis cultivar)
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Sapindales
Family: Rutaceae
Subfamily: Aurantioideae
Genus: Citrus
L.
Species and hybrids

Ancestral species:
Citrus maxima – Pomelo
Citrus medica – Citron
Citrus reticulata – Mandarin orange
Citrus micrantha – a papeda
Citrus hystrix – Kaffir lime
Citrus cavaleriei – Ichang papeda
Citrus japonica – Kumquat

Important hybrids:
Citrus × aurantiifolia – Key lime
Citrus × aurantium – Bitter orange
Citrus × latifolia – Persian lime
Citrus × limon – Lemon
Citrus × limonia – Rangpur
Citrus × paradisi – Grapefruit
Citrus × sinensis – Sweet orange
Citrus × tangerina – Tangerine
See also below for other species and hybrids.

Synonyms[2]
  • Aurantium Mill.
  • Citreum Mill.
  • ×Citrofortunella J.W.Ingram & H.E.Moore
  • ×Citroncirus J.W.Ingram & H.E.Moore
  • Citrophorum Neck.
  • Eremocitrus Swingle
  • Feroniella Swingle
  • Fortunella Swingle
  • Limon Mill.
  • Microcitrus Swingle
  • Oxanthera Montrouz.
  • Papeda Hassk.
  • Pleurocitrus Tanaka
  • Poncirus Raf.
  • Pseudaegle Miq.
  • Sarcodactilis C.F.Gaertn.

Citrus is a genus of flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes.

Citrus is native to South Asia, East Asia, Southeast Asia, Melanesia, and Australia. Indigenous people in these areas have used and domesticated various species since ancient times. Its cultivation first spread into Micronesia and Polynesia through the Austronesian expansion (c. 3000–1500 BCE). Later, it was spread to the Middle East and the Mediterranean (c. 1200 BCE) via the incense trade route, and from Europe to the Americas.

Renowned for their highly fragrant aromas and complex flavor, citrus are among the most popular fruits in cultivation. With a propensity to hybridize between species, making their taxonomy complicated, there are numerous varieties encompassing a wide range of appearance and fruit flavors.

Evolution

Evolutionary history

The large citrus fruit of today evolved originally from small, edible berries over millions of years. Citrus species began to diverge from a common ancestor about 15 million years ago, at about the same time that Severinia (such as the Chinese box orange) diverged from the same ancestor. About 7 million years ago, the ancestors of Citrus split into the main genus, Citrus, and the genus Poncirus (such as the trifoliate orange), which is closely enough related that it can still be hybridized with all other citrus and used as rootstock. These estimates are made using genetic mapping of plant chloroplasts.[3] A DNA study published in Nature in 2018 concludes that the genus Citrus evolved in the foothills of the Himalayas, in the area of Assam (India), western Yunnan (China), and northern Myanmar.[4]

Map of inferred original wild ranges of the main Citrus cultivars, and selected relevant wild taxa[5]

The three ancestral species in the genus Citrus associated with modern Citrus cultivars are the mandarin orange, pomelo, and citron. Almost all of the common commercially important citrus fruits (sweet oranges, lemons, grapefruit, limes, and so on) are hybrids between these three species, their main progenies, and other wild Citrus species within the last few thousand years.[6][7][8]

Citrus plants are native to subtropical and tropical regions of Asia, Island Southeast Asia, Near Oceania, and northeastern and central Australia. Domestication of citrus species involved much hybridization and introgression, leaving much uncertainty about when and where domestication first happened.[6] A genomic, phylogenic, and biogeographical analysis by Wu et al. (2018) has shown that the center of origin of the genus Citrus is likely the southeast foothills of the Himalayas, in a region stretching from eastern Assam, northern Myanmar, to western Yunnan. It diverged from a common ancestor with Poncirus trifoliata. A change in climate conditions during the Late Miocene (11.63 to 5.33 mya) resulted in a sudden speciation event. The species resulting from this event include the citrons (Citrus medica) of South Asia; the pomelos (C. maxima) of Mainland Southeast Asia; the mandarins (C. reticulata), kumquats (C. japonica), mangshanyegan (C. mangshanensis), and ichang papedas (C. cavaleriei) of southeastern China; the kaffir limes (C. hystrix) of Island Southeast Asia; and the biasong and samuyao (C. micrantha) of the Philippines.[6][5]

This was followed by the spread of citrus species into Taiwan and Japan in the Early Pliocene (5.33 to 3.6 mya), resulting in the tachibana orange (C. tachibana); and beyond the Wallace Line into Papua New Guinea and Australia during the Early Pleistocene (2.5 million to 800,000 years ago), where further speciation events created in the Australian limes.[6][5]

Fossil record

A fossil leaf from the Pliocene of Valdarno (Italy) is described as †Citrus meletensis.[9] In China, fossil leaf specimens of †Citrus linczangensis have been collected from coal-bearing strata of the Bangmai Formation in the Bangmai village, about 10 km (6 miles) northwest of Lincang City, Yunnan. The Bangmai Formation contains abundant fossil plants and is considered to be of late Miocene age. Citrus linczangensis and C. meletensis share some important characters, such as an intramarginal vein, an entire margin, and an articulated and distinctly winged petiole.[10]

Taxonomy

Main article: Citrus taxonomy

Many cultivated Citrus species are natural or artificial hybrids of a small number of core ancestral species, including the citron, pomelo, and mandarin. Natural and cultivated citrus hybrids include commercially important fruit such as oranges, grapefruit, lemons, limes, and some tangerines. The multiple hybridisations have made the taxonomy of Citrus complex.[11][12]

Many Citrus species are hybrids of citron, mandarin and pomelo.[12] There are also kumquat and trifoliate orange hybrids.

Apart from these core species, Australian limes and the recently discovered mangshanyegan are grown. Kumquats and Clymenia spp. are now generally considered to belong within the genus Citrus.[13] The false oranges, Oxanthera from New Caledonia, have been transferred to the Citrus genus on phylogenetic evidence.[14][15]

History

Domestication

The earliest introductions of citrus species by human migrations was during the Austronesian expansion (c. 3000–1500 BCE), where Citrus hystrix, Citrus macroptera, and Citrus maxima were among the canoe plants carried by Austronesian voyagers eastwards into Micronesia and Polynesia.[16]

The citron (Citrus medica) was also introduced early into the Mediterranean basin from India and Southeast Asia. It was introduced via two ancient trade routes: an overland route through Persia, the Levant and the Mediterranean islands; and a maritime route through the Arabian Peninsula and Ptolemaic Egypt into North Africa. Although the exact date of the original introduction is unknown due to the sparseness of archaeobotanical remains, the earliest evidence are seeds recovered from the Hala Sultan Tekke site of Cyprus, dated to around 1200 BCE. Other archaeobotanical evidence includes pollen from Carthage dating back to the 4th century BCE; and carbonized seeds from Pompeii dated to around the 3rd to 2nd century BCE. The earliest complete description of the citron was written by Theophrastus, c. 310 BCE.[17][18][19]

Lemons, pomelos, and sour oranges were introduced to the Mediterranean by Arab traders around the 10th century CE. Sweet oranges were brought to Europe by the Genoese and Portuguese from Asia during the 15th to 16th century. Mandarins were not introduced until the 19th century.[17][18][19]

Oranges were introduced to Florida by Spanish colonists.[20][21] In cooler parts of Europe, citrus fruit was grown in orangeries starting in the 17th century; many were as much status symbols as functional agricultural structures.[22]

Etymology

The generic name originated from Latin, where it referred to either the citron (C. medica) or a conifer tree (Thuja). The Latin word is related to the ancient Greek word for cedar, κέδρος (kédros). This may be due to perceived similarities in the smell of citrus leaves and fruit with that of cedar.[23]

Description

Tree

Citrus plants are large shrubs or small to moderate-sized trees, reaching 5–15 m (16–49 ft) tall, with spiny shoots and alternately arranged evergreen leaves with an entire margin.[24] The flowers are solitary or in small corymbs, each flower 2–4 cm (0.79–1.57 in) diameter, with five (rarely four) white petals and numerous stamens; they are often very strongly scented, due to the presence of essential oil glands.[25]

Fruit

For a more comprehensive list, see List of citrus fruits.
Structure of the botanical hesperidium

The fruit is a hesperidium, a specialised berry with multiple carpels, globose to elongated,[25][26] 4–30 cm (1.6–11.8 in) long and 4–20 cm (1.6–7.9 in) diameter, with a leathery rind or "peel" called a pericarp. The outermost layer of the pericarp is an "exocarp" called the flavedo, commonly referred to as the zest. The middle layer of the pericarp is the mesocarp, which in citrus fruits consists of the white, spongy "albedo", or "pith". The innermost layer of the pericarp is the endocarp. This surrounds a variable number of carpels, shaped as radial segments. The seeds, if present, develop inside the carpels. The space inside each segment is a locule filled with juice vesicles, or "pulp". From the endocarp, string-like "hairs" extend into the locules, which provide nourishment to the fruit as it develops.[25][27][28] The genus is commercially important with cultivars of many species grown for their fruit. Some cultivars have been developed to be easy to peel and seedless, meaning they are parthenocarpic.[26]

The fragrance of citrus fruits is conferred by flavonoids and limonoids in the rind. The flavonoids include various flavanones and flavones.[29] The carpels are juicy; they contain a high quantity of citric acid, which with other organic acids including ascorbic acid (vitamin C) give them their characteristic sharp taste.[30] Citrus fruits are diverse in size and shape, as well as in color and flavor, reflecting their biochemistry;[31][32] for instance, grapefruit is made bitter-tasting by a flavanone, naringin.[30]


Cultivation

Further information: Citrus production
Mediterranean Mandarin (Citrus × deliciosa) plantation, Mallorca

Citrus trees hybridise very readily – depending on the pollen source, plants grown from a Persian lime's seeds can produce fruit similar to grapefruit. Most commercial citrus cultivation uses trees produced by grafting the desired fruiting cultivars onto rootstocks selected for disease resistance and hardiness.

The colour of citrus fruits only develops in climates with a (diurnal) cool winter. In tropical regions with no winter at all, citrus fruits remain green until maturity, hence the tropical "green oranges".[33]

The terms 'ripe' and 'mature' are widely used synonymously, but they mean different things. A mature fruit is one that has completed its growth phase. Ripening is the sequence of changes within the fruit from maturity to the beginning of decay. These changes involve the conversion of starches to sugars, a decrease in acids, softening, and s change in the fruit's colour.[34]

Citrus fruits are non-climacteric and respiration slowly declines and the production and release of ethylene is gradual.[35] The fruits do not go through a ripening process in the sense that they become "tree ripe". Some fruits, for example cherries, physically mature and then continue to ripen on the tree. Other fruits, such as pears, are picked when mature, but before they ripen, then continue to ripen off the tree. Citrus fruits pass from immaturity to maturity to overmaturity while still on the tree. Once they are separated from the tree, they do not increase in sweetness or continue to ripen, but proceed to decay.

With oranges, colour cannot be used as an indicator of ripeness because sometimes the rinds turn orange long before the oranges are ready to eat. Tasting is the only way to know whether they are ripe.

Citrus trees are not generally frost hardy. Mandarin oranges (C. reticulata) tend to be the hardiest of the common Citrus species and can withstand short periods down to as cold as −10 °C (14 °F), but realistically temperatures not falling below −2 °C (28 °F) are required for successful cultivation. Tangerines, tangors and yuzu can be grown outside even in regions with more marked subfreezing temperatures in winter, although this may affect fruit quality. A few hardy hybrids can withstand temperatures well below freezing, but do not produce quality fruit. Lemons can be commercially grown in cooler-summer/moderate-winter, coastal Southern California, because sweetness is neither attained nor expected in retail lemon fruit. The related trifoliate orange (C. trifoliata) can survive below −20 °C (−4 °F); its fruit are astringent and inedible unless cooked, but a few better-tasting cultivars and hybrids have been developed (see citranges).

The trees thrive in a consistently sunny, humid environment with fertile soil and adequate rainfall or irrigation. Abandoned trees in valleys may suffer, yet survive, the dry summer of Central California's Inner Coast Ranges. At any age, citrus grows well enough with infrequent irrigation in partial shade, but the fruit crop is smaller. Being of tropical and subtropical origin, oranges, like all citrus, are broadleaved and evergreen. They do not drop leaves except when stressed. The stems of many varieties have large sharp thorns. The trees flower in the spring, and fruit is set shortly afterward. Fruit begins to ripen in fall or early winter, depending on cultivar, and develops increasing sweetness afterward. Some cultivars of tangerines ripen by winter. Some, such as the grapefruit, take up to 18 months to ripen.

Production

Further information: Citrus production
Major producer regions

According to the UN Food and Agriculture Organization, world production of all citrus fruits in 2016 was 124 million metric tons (122,000,000 long tons; 137,000,000 short tons), with about half of this production as oranges.[36] At US $15.2 billion equivalent in 2018, citrus trade[37] makes up nearly half of the world fruit trade, which was US$32.1 billion for the same year.[38] According to the United Nations Conference on Trade and Development (UNCTAD), citrus production grew during the early 21st century mainly by the increase in cultivation areas, improvements in transportation and packaging, rising incomes and consumer preference for healthy foods.[36] In 2019–20, world production of oranges was estimated to be 47.5 million metric tons (46,700,000 long tons; 52,400,000 short tons), led by Brazil, Mexico, the European Union, and China as the largest producers.[39]

Pests and diseases

For a more comprehensive list, see List of citrus diseases.

Among the diseases of citrus plantations are citrus black spot (a fungus), citrus canker (a bacterium), citrus greening (a bacterium, spread by an insect pest), and sweet orange scab (a fungus, Elsinöe australis).[40] Citrus plants are liable to infestation by ectoparasites which act as vectors to plant diseases: for example, aphids transmit the damaging citrus tristeza virus,[41] while the aphid-like Asian citrus psyllid can carry the bacterium which causes the serious citrus greening disease.[42][43] This threatens production in Florida[44][45] and California.[43]

Deficiency diseases

Citrus plants can develop the deficiency condition chlorosis, characterized by yellowing leaves.[46] The condition is often caused by an excessively high pH (alkaline soil), which prevents the plant from absorbing nutrients such as iron, magnesium, and zinc needed to produce chlorophyll.[47]

Uses

Culinary

Wikibooks Cookbook has a recipe/module on

Many citrus fruits, such as oranges, tangerines, grapefruits, and clementines, are generally eaten fresh.[26] They are typically peeled and can be easily split into segments.[26] Grapefruit is more commonly halved and eaten out of the skin with a spoon.[48] Special spoons (grapefruit spoons) with serrated tips are designed for this purpose. Orange and grapefruit juices are also popular breakfast beverages. More acidic citrus, such as lemons and limes, are generally not eaten on their own. Meyer lemons can be eaten out of hand with the fragrant skin; they are both sweet and sour. Lemonade or limeade are popular beverages prepared by diluting the juices of these fruits and adding sugar. Lemons and limes are also used in cooked dishes, or sliced and used as garnishes. Their juice is used as an ingredient in a variety of dishes; it can commonly be found in salad dressings and squeezed over cooked fish, meat, or vegetables.

A variety of flavours can be derived from different parts and treatments of citrus fruits.[26] The rind and oil of the fruit is generally bitter, especially when cooked, so is often combined with sugar. The fruit pulp can vary from sweet to extremely sour. Marmalade, a condiment derived from cooked orange or lemon, can be especially bitter, but is usually sweetened with sugar to cut the bitterness and produce a jam-like result. Lemon or lime is commonly used as a garnish for water, soft drinks, or cocktails. Citrus juices, rinds, or slices are used in a variety of mixed drinks. The colourful outer skin of some citrus fruits, known as zest, is used as a flavouring in cooking; the whole of the bitter orange is cooked with sugar to make marmalade.

As ornamental plants

The Versailles Orangerie, 1686

By the 17th century, orangeries were added to great houses in Europe, both to enable the fruit to be grown locally and for prestige, as seen in the Versailles Orangerie completed in 1686.[49] Some modern hobbyists grow dwarf citrus in containers or greenhouses in areas where the weather is too cold to grow it outdoors; Citrofortunella hybrids have good cold resistance..[50]

Effects on humans

Some Citrus species contain significant amounts of furanocoumarins.[51][52] In humans, some of these act as strong photosensitizers when applied topically to the skin, while others interact with medications when taken orally in the grapefruit juice effect.[51]

Due to the photosensitizing effects of certain furanocoumarins, some Citrus species cause phytophotodermatitis,[53] a potentially severe skin inflammation resulting from contact with a light-sensitizing botanical agent followed by exposure to ultraviolet light. In Citrus species, the primary photosensitizing agent appears to be bergapten,[54] a linear furanocoumarin derived from psoralen. This claim has been confirmed for lime[55][56] and bergamot. In particular, bergamot essential oil has a higher concentration of bergapten (3000–3600& mg/kg) than any other Citrus-based essential oil.[57]

A systematic review indicates that citrus fruit consumption is associated with a 10% reduction of risk for developing breast cancer.[58]

In art and culture

Giovanna Garzoni's Still Life with Bowl of Citrons, late 1640s

Lemons appear in paintings, pop art, and novels.[59] A wall painting in the tomb of Nakht in 15th century BC Egypt depicts a woman in a festival, holding a lemon. In the 17th century, Giovanna Garzoni painted a Still Life with Bowl of Citrons, the fruits still attached to leafy flowering twigs, with a wasp on one of the fruits. The impressionist Edouard Manet depicted a lemon on a pewter plate. In modern art, Arshile Gorky painted Still Life with Lemons in the 1930s.[59]

Citrus fruits "were the clear status symbols of the nobility in the ancient Mediterranean", according to the paleoethnobotanist Dafna Langgut.[60] In Louisa May Alcott's 1868 novel Little Women, the character Amy March states that "It's nothing but limes now, for everyone is sucking them in their desks in schooltime, and trading them off for pencils, bead rings, paper dolls, or something else… If one girl likes another, she gives her a lime; if she’s mad with her, she eats one before her face, and doesn’t offer even a suck."[60]

In 20th century American self-improvement culture, Dale Carnegie advised readers "If You Have a Lemon, Make a Lemonade", meaning to make the best of what you have.[60] In the 21st century, a defective machine such as a car is called a lemon.[60]

See also

References

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