A mango is an edible stone fruit produced by the tropical tree Mangifera indica. It originated from the region between northwestern Myanmar, Bangladesh, and northeastern India.[1][2]M. indica has been cultivated in South and Southeast Asia since ancient times resulting in two types of modern mango cultivars: the “Indian type” and the “Southeast Asian type”.[1][2] Other species in the genus Mangifera also produce edible fruits that are also called “mangoes”, the majority of which are found in the Malesian ecoregion.[3]
Worldwide, there are several hundred cultivars of mango. Depending on the cultivar, mango fruit varies in size, shape, sweetness, skin color, and flesh color, which may be pale yellow, gold, green, or orange.[4] Mango is the national fruit of India, Pakistan and the Philippines,[5][6] while the mango tree is the national tree of Bangladesh.[7]
Etymology
The English word mango (plural mangoes or mangos) originated in the 16th century from the Portuguese word manga, from the Malaymangga, and ultimately from the Tamilmā (மா, ‘mango tree’) + kāy (ங்காய், ‘unripe fruit/vegetable’)[8][9][10] or the Malayalammāṅṅa (മാവ്, ‘mango tree’) + kāya (കായ, ‘unripe fruit’).[11] The scientific name, Mangifera indica, refers to a plant bearing mangoes in India.[10]
Description
Mango trees grow to 30–40 metres (98–131 feet) tall, with a crown radius of 10–15 m (33–49 ft). The trees are long-lived, as some specimens still fruit after 300 years.[12]
In deep soil, the taproot descends to a depth of 6 m (20 ft), with profuse, wide-spreading feeder roots and anchor roots penetrating deeply into the soil.[4] The leaves are evergreen, alternate, simple, 15–35 centimetres (6–14 inches) long, and 6–16 cm (2+1⁄2–6+1⁄2 in) broad; when the leaves are young they are orange-pink, rapidly changing to a dark, glossy red, then dark green as they mature.[4] The flowers are produced in terminal panicles 10–40 cm (4–15+1⁄2 in) long; each flower is small and white with five petals 5–10 millimetres (3⁄16–3⁄8 in) long, with a mild, sweet fragrance.[4] Over 500 varieties of mangoes are known,[4] many of which ripen in summer, while some give a double crop.[13] The fruit takes four to five months from flowering to ripening.[4]
The ripe fruit varies according to cultivar in size, shape, color, sweetness, and eating quality.[4] Depending on the cultivar, fruits are variously yellow, orange, red, or green.[4] The fruit has a single flat, oblong pit that can be fibrous or hairy on the surface and does not separate easily from the pulp.[4] The fruits may be somewhat round, oval, or kidney-shaped, ranging from 5–25 centimetres (2–10 in) in length and from 140 grams (5 oz) to 2 kilograms (5 lb) in weight per individual fruit.[4] The skin is leather-like, waxy, smooth, and fragrant, with colors ranging from green to yellow, yellow-orange, yellow-red, or blushed with various shades of red, purple, pink, or yellow when fully ripe.[4]
Ripe intact mangoes give off a distinctive resinous, sweet smell.[4] Inside the pit 1–2 mm (0.039–0.079 in) thick is a thin lining covering a single seed, 4–7 cm (1.6–2.8 in) long. Mangoes have recalcitrant seeds which do not survive freezing and drying.[14] Mango trees grow readily from seeds, with germination success highest when seeds are obtained from mature fruits.[4]
From their center of origin, mangoes diverged into two genetically distinct populations: the subtropical Indian group and the tropical Southeast Asian group. The Indian group is characterized by having monoembryonic fruits, while polyembryonic fruits characterize the Southeast Asian group.[1][2]
It was previously believed that mangoes originated from a single domestication event in South Asia before being spread to Southeast Asia, but a 2019 study found no evidence of a center of diversity in India. Instead, it identified a higher unique genetic diversity in Southeast Asian cultivars than in Indian cultivars, indicating that mangoes may have originally been domesticated first in Southeast Asia before being introduced to South Asia. However, the authors also cautioned that the diversity in Southeast Asian mangoes might be the result of other reasons (like interspecific hybridization with other Mangifera species native to the Malesian ecoregion). Nevertheless, the existence of two distinct genetic populations also identified by the study indicates that the domestication of the mango is more complex than previously assumed and would at least indicate multiple domestication events in Southeast Asia and South Asia.[1][2]
There are hundreds of named mango cultivars. In mango orchards, several cultivars are often grown to improve pollination. Many desired cultivars are monoembryonic and must be propagated by grafting, or they do not breed true. A common monoembryonic cultivar is ‘Alphonso’, an important export product, considered “the king of mangoes”.[16]
Cultivars that excel in one climate may fail elsewhere. For example, Indian cultivars such as ‘Julie,’ a prolific cultivar in Jamaica, require annual fungicide treatments to escape the lethal fungal diseaseanthracnose in Florida. Asian mangoes are resistant to anthracnose.[17]
The current world market is dominated by the cultivar ‘Tommy Atkins’, a seedling of ‘Haden’ that first fruited in 1940 in southern Florida and was initially rejected commercially by Florida researchers.[18] Growers and importers worldwide have embraced the cultivar for its excellent productivity and disease resistance, shelf life, transportability, size, and appealing color.[19] Although the Tommy Atkins cultivar is commercially successful, other cultivars may be preferred by consumers for eating pleasure, such as Alphonso.[16][19]
Generally, ripe mangoes have an orange-yellow or reddish peel and are juicy for eating, while exported fruit are often picked while underripe with green peels. Although producing ethylene while ripening, unripened exported mangoes do not have the same juiciness or flavor as fresh fruit.
Distribution and habitat
Mango tree in Palestine
From tropical Asia, mangoes were introduced to East Africa by Arab and Persian traders in the ninth to tenth centuries.[20] The 14th-century Moroccan traveler Ibn Battuta reported it at Mogadishu.[21] It was spread further into other areas around the world during the Colonial Era. The Portuguese Empire spread the mango from their colony in Goa to East and West Africa. From West Africa, they introduced it to Brazil from the 16th to the 17th centuries. From Brazil, it spread northwards to the Caribbean and eastern Mexico by the mid to late 18th century. The Spanish Empire also introduced mangoes directly from the Philippines to western Mexico via the Manila galleons from at least the 16th century. Mangoes were only introduced to Florida by 1833.[2][22]
Cultivation
The mango is now cultivated in most frost-free tropical and warmer subtropical climates. It is cultivated extensively in South Asia, Southeast Asia, East and West Africa, the tropical and subtropical Americas, and the Caribbean.[23] Mangoes are also grown in Andalusia, Spain (mainly in Málaga province), as its coastal subtropical climate is one of the few places in mainland Europe that permits the growth of tropical plants and fruit trees. The Canary Islands are another notable Spanish producer of the fruit. Other minor cultivators include North America (in South Florida and the California Coachella Valley), Hawai’i, and Australia.[24]
Many commercial cultivars are grafted onto the cold-hardy rootstock of the Gomera-1 mango cultivar, originally from Cuba. Its root system is well adapted to a coastal Mediterranean climate.[25] Many of the 1,000+ mango cultivars are easily cultivated using grafted saplings, ranging from the “turpentine mango” (named for its strong taste of turpentine[26]) to the Bullock’s Heart. Dwarf or semidwarf varieties serve as ornamental plants and can be grown in containers. A wide variety of diseases can afflict mangoes.[citation needed]
*includes mangosteens and guavas. Source: FAOSTAT of the United Nations[27]
A breakthrough in mango cultivation was the use of potassium nitrate and ethrel to induce flowering in mangoes. The discovery was made by FilipinohorticulturistRamon Barba in 1974 and was developed from the unique traditional method of inducing mango flowering using smoke in the Philippines. It allowed mango plantations to induce regular flowering and fruiting year-round. Previously, mangoes were seasonal because they only flowered every 16 to 18 months. The method is now used in most mango-producing countries.[28][29]
Production
In 2022, world production of mangoes (report includes mangosteens and guavas) was 59 million tonnes, led by India with 44% of the total (table).
Mangoes are generally sweet, although the taste and texture of the flesh vary across cultivars; some, such as Alphonso, have a soft, pulpy, juicy texture similar to an overripe plum, while others, such as Tommy Atkins, are firmer with a fibrous texture.[30]
The skin of unripe, pickled, or cooked mango can be eaten, but it has the potential to cause contact dermatitis of the lips, gingiva, or tongue in susceptible people.[31]
Mangoes are used in many cuisines. Sour, unripe mangoes are used in chutneys (i.e., mango chutney), pickles, daals and other side dishes in Indian cuisine. A summer drink called aam panna is made with mangoes. Mango pulp made into jelly or cooked with red gram dhal and green chilies may be served with cooked rice. Mango lassi is consumed throughout South Asia, prepared by mixing ripe mangoes or mango pulp with buttermilk and sugar. Ripe mangoes are also used to make curries. Aamras is a thick juice made of mangoes with sugar or milk and is consumed with chapatis or pooris. The pulp from ripe mangoes is also used to make jam called mangada. Andhra aavakaaya is a pickle made from raw, unripe, pulpy, and sour mango mixed with chili powder, fenugreek seeds, mustard powder, salt, and groundnut oil. Mango is also used to make dahl and chunda (a sweet and spicy, grated mango delicacy). In Indonesian cuisine, unripe mango is processed into asinan, rujak and sambal pencit/mangga muda,[32] or eaten with edible salt.[33]
Mangoes are used to make murabba (fruit preserves), muramba (a sweet, grated mango delicacy), amchur (dried and powdered unripe mango), and pickles, including a spicy mustard-oil pickle and alcohol. Ripe mangoes are cut into thin layers, desiccated, folded, and then cut. The fruit is also added to cereal products such as muesli and oat granola.
Pieces of mango can be mashed and used as a topping on ice cream or blended with milk and ice as milkshakes. Sweet glutinous rice is flavored with coconut, then served with sliced mango as mango sticky rice. In other parts of Southeast Asia, mangoes are pickled with fish sauce and rice vinegar. Green mangoes can be used in mango salad with fish sauce and dried shrimp. Mango with condensed milk may be used as a topping for shaved ice.
Raw green mangoes can be sliced and eaten like a salad.[34] In most parts of Southeast Asia, they are commonly eaten with fish sauce, vinegar, soy sauce, or with a dash of salt (plain or spicy) – a combination usually known as “mango salad” in English.[35]
Major flavor chemicals of ‘Alphonso’ mango from India
In the Philippines, green mangoes are also commonly eaten with bagoong (salty fish or shrimp paste), salt, soy sauce, vinegar, or chilis.[36][37]Mango float and mango cake, which use slices of ripe mangoes, are eaten in the Philippines.[38][39]Dried strips of sweet, ripe mango (sometimes combined with seedless tamarind to form mangorind) are also consumed. Mangoes may be used to make juices, mango nectar, and as a flavoring and major ingredient in mango ice cream and sorbetes.
Phytochemical and nutrient content appears to vary across mango cultivars.[48] Up to 25 different carotenoids have been isolated from mango pulp, the densest of which was beta-carotene, which accounts for the yellow-orange pigmentation of most mango cultivars.[49] Mango leaves also have significant polyphenol content, including xanthonoids, mangiferin and gallic acid.[50]
Flavor
The flavor of mango fruits is conferred by several volatile organic chemicals mainly belonging to terpene, furanone, lactone, and ester classes. Different varieties or cultivars of mangoes can have flavors made up of different volatile chemicals or the same volatile chemicals in different quantities.[51] In general, New World mango cultivars are characterized by the dominance of δ-3-carene, a monoterpene flavorant; whereas, high concentration of other monoterpenes such as (Z)-ocimene and myrcene, as well as the presence of lactones and furanones, is the unique feature of Old World cultivars.[52][53][54] In India, ‘Alphonso’ is one of the most popular cultivars. In ‘Alphonso’ mango, the lactones, and furanones are synthesized during ripening, whereas terpenes and the other flavorants are present in both the developing (immature) and ripening fruits.[55][56][57]Ethylene, a ripening-related hormone well known to be involved in ripening of mango fruits, causes changes in the flavor composition of mango fruits upon exogenous application, as well.[58][59] In contrast to the huge amount of information available on the chemical composition of mango flavor, the biosynthesis of these chemicals has not been studied in depth; only a handful of genes encoding the enzymes of flavor biosynthetic pathways have been characterized to date.[60][61][62][63]
Toxicity
Contact with oils in mango leaves, stems, sap, and skin can cause dermatitis and anaphylaxis in susceptible individuals.[4][31][64] Those with a history of contact dermatitis induced by urushiol (an allergen found in poison ivy, poison oak, or poison sumac) may be most at risk for mango contact dermatitis.[65] Other mango compounds potentially responsible for dermatitis or allergic reactions include mangiferin.[4] Cross-reactions may occur between mango allergens and urushiol.[66] Sensitized individuals may not be able to eat peeled mangos or drink mango juice safely.[4]
When mango trees are flowering in spring, local people with allergies may experience breathing difficulty, itching of the eyes, or facial swelling, even before flower pollen becomes airborne.[4] In this case, the irritant is likely to be the vaporizedessential oil from flowers.[4] During the primary ripening season of mangoes, contact with mango plant parts – primarily sap, leaves, and fruit skin[4] – is the most common cause of plant dermatitis in Hawaii.[67]
Nutrition
A raw mango is 84% water, 15% carbohydrates, 1% protein, and has negligible fat (table). In a reference amount of 100 g (3.5 oz), raw mango supplies 60 calories and is a rich source of vitamin C (44% of the Daily Value, DV) with moderate amounts of folate (11% DV) and copper (12% DV), while other micronutrients are low in content (table).
The mango is the national fruit of India.[68][69] It is also the national tree of Bangladesh.[70][71] In India, harvest and sale of mangoes is during March–May and this is annually covered by news agencies.[16]
The mango has a traditional context in the culture of South Asia. In his edicts, the Mauryan emperor Ashoka references the planting of fruit- and shade-bearing trees along imperial roads:
“On the roads banyan-trees were caused to be planted by me, (in order that) they might afford shade to cattle and men, (and) mango-groves were caused to be planted.”
In medieval India, the Indo-Persian poet Amir Khusrau termed the mango “Naghza Tarin Mewa Hindustan” – “the fairest fruit of Hindustan”. Mangoes were enjoyed at the court of the Delhi Sultan Alauddin Khijli. The Mughal Empire was especially fond of the fruits: Babur praises the mango in his Babarnameh. At the same time, Sher Shah Suri inaugurated the creation of the Chaunsa variety after his victory over the Mughal emperor Humayun. Mughal patronage of horticulture led to the grafting of thousands of mangoes varieties, including the famous Totapuri, which was the first variety to be exported to Iran and Central Asia.[citation needed]Akbar (1556–1605) is said to have planted a mango orchard of 100,000 trees near Darbhanga, Bihar,[72] while Jahangir and Shah Jahan ordered the planting of mango orchards in Lahore and Delhi and the creation of mango-based desserts.[73]
The Jain goddess Ambika is traditionally represented as sitting under a mango tree.[74] Mango blossoms are also used in the worship of the goddess Saraswati. Mango leaves decorate archways and doors in Indian houses during weddings and celebrations such as Ganesh Chaturthi. Mango motifs and paisleys are widely used in different Indian embroidery styles, and are found in Kashmiri shawls, Kanchipuram and silk sarees. In Tamil Nadu, the mango is referred to as one of the three royal fruits, along with banana and jackfruit, for their sweetness and flavor.[75] This triad of fruits is referred to as ma-pala-vazhai. The classical Sanskrit poet Kalidasa sang the praises of mangoes.[76]
Apples grown from seed tend to be very different from those of their parents, and the resultant fruit frequently lacks desired characteristics. For commercial purposes, including botanical evaluation, apple cultivars are propagated by clonal grafting onto rootstocks. Apple trees grown without rootstocks tend to be larger and much slower to fruit after planting. Rootstocks are used to control the speed of growth and the size of the resulting tree, allowing for easier harvesting.
There are more than 7,500 cultivars of apples. Different cultivars are bred for various tastes and uses, including cooking, eating raw, and cider or apple juice production. Trees and fruit are prone to fungal, bacterial, and pest problems, which can be controlled by a number of organic and non-organic means. In 2010, the fruit’s genome was sequenced as part of research on disease control and selective breeding in apple production.
Etymology
The word apple, whose Old English ancestor is æppel, is descended from the Proto-Germanic noun *aplaz, descended in turn from Proto-Indo-European *h₂ébōl.[3] As late as the 17th century, the word also functioned as a generic term for all fruit, including nuts. This can be compared to the 14th-century Middle English expression appel of paradis, meaning a banana.[4]
Description
The apple is a deciduous tree, generally standing 2 to 4.5 metres (6 to 15 feet) tall in cultivation and up to 15 m (49 ft) in the wild, though more typically 2 to 10 m (6.5 to 33 ft).[5][1] When cultivated, the size, shape and branch density are determined by rootstock selection and trimming method.[5] Apple trees may naturally have a rounded to erect crown with a dense canopy of leaves.[6] The bark of the trunk is dark gray or gray-brown, but young branches are reddish or dark-brown with a smooth texture.[1][7] Young twigs are covered in fine downy hairs; they become hairless when older.[7]
The buds are egg-shaped and dark red or purple in color; they range in size from 3 to 5 millimeters, but are usually less than 4 mm. The bud scales have very hairy edges. When emerging from the buds, the leaves are convolute, meaning that their edges overlap each other.[1] Leaves can be simple ovals (elliptic), medium or wide in width, somewhat egg-shaped with the wider portion toward their base (ovate), or even with sides that are more parallel to each other instead of curved (oblong) with a narrow pointed end.[7][1] The edges have broadly-angled teeth, but do not have lobes. The top surface of the leaves are glabrescent, almost hairless, while the undersides are densely covered in fine hairs.[1] The leaves are attached alternately by short leaf stems 1-to-3.5 cm (1⁄2-to-1+1⁄2 in) long.[6][1]
Blossoms are produced in spring simultaneously with the budding of the leaves and are produced on spurs and some long shoots.[5] When the flower buds first begin to open the petals are rose-pink and fade to white or light pink when fully open with each flower 3-to-4-centimeter (1-to-1+1⁄2-inch) in diameter.[1] The five-petaled flowers are group in an inflorescence consisting of a cyme with 3–7 flowers.[8] The central flower of the inflorescence is called the “king bloom”; it opens first and can develop a larger fruit.[6] Open apple blossoms are damaged by even brief exposures to temperatures −2 °C (28 °F) or less, although the overwintering wood and buds are hardy down to −40 °C (−40 °F).[8]
Apple blossoms
Botanical illustration
Fruit
The fruit is a pome that matures in late summer or autumn.[1] The true fruits or carpels are the harder interior chambers inside the apple’s core. There are usually five carpels inside an apple, but there may be as few as three. Each of the chambers contains one or two seeds.[9] The edible flesh is formed from the receptacle at the base of the flower.[10]
How apple fruit derives from flower structures
The seeds are egg- to pear-shaped and may be colored from light brown or tan to a very dark brown, often with red shades or even purplish-black. They may have a blunt or sharp point.[11] The five sepals remain attached and stand out from the surface of the apple.[1]
The size of the fruit varies widely between cultivars, but generally has a diameter between 2.5 and 12 cm (1 and 5 in).[7] The shape is quite variable and may be nearly round, elongated, conical, or short and wide.[12]
The groundcolor of ripe apples is yellow, green, yellow-green or whitish yellow. The overcolor of ripe apples can be orange-red, pink-red, red, purple-red or brown-red. The overcolor amount can be 0–100%.[13] The skin may be wholly or partly russeted, making it rough and brown. The skin is covered in a protective layer of epicuticular wax.[14] The skin may also be marked with scattered dots.[1] The flesh is generally pale yellowish-white, though it can be pink, yellow or green.[13]
Apples can have any amount of overcolor, a darker tint over a pale groundcolor.
The apple as a species has more than 100 alternative scientific names, or synonyms.[17] In modern times, Malus pumila and Malus domestica are the two main names in use. M. pumila is the older name, but M. domestica has become much more commonly used starting in the 21st century, especially in the western world. Two proposals were made to make M. domestica a conserved name: the earlier proposal was voted down by the Committee for Vascular Plants of the IAPT in 2014, but in April 2017 the Committee decided, with a narrow majority, that the newly popular name should be conserved.[18] The General Committee of the IAPT decided in June 2017 to approve this change, officially conserving M. domestica.[19] Nevertheless, some works published after 2017 still use M. pumila as the correct name, under an alternate taxonomy.[2]
When first classified by Linnaeus in 1753, the pears, apples, and quinces were combined into one genus that he named Pyrus and he named the apple as Pyrus malus. This was widely accepted, however the botanist Philip Miller published an alternate classification in The Gardeners Dictionary with the apple species separated from Pyrus in 1754. He did not clearly indicate that by Malus pumila he meant the domesticated apple. Nonetheless, it was used as such by many botanists. When Moritz Balthasar Borkhausen published his scientific description of the apple in 1803 it may have been a new combination of P. malus var. domestica, but this was not directly referenced by Borkhausen.[17] The earliest use of var. domestica for the apple was by Georg Adolf Suckow in 1786.[2]
Apples are diploid, with two sets of chromosomes per cell (though triploid cultivars, with three sets, are not uncommon), have 17 chromosomes and an estimated genome size of approximately 650 Mb. Several whole genome sequences have been completed and made available. The first one in 2010 was based on the diploid cultivar ‘Golden Delicious‘.[20] However, this first whole genome sequence contained several errors,[21] in part owing to the high degree of heterozygosity in diploid apples which, in combination with an ancient genome duplication, complicated the assembly. Recently, double- and trihaploid individuals have been sequenced, yielding whole genome sequences of higher quality.[22][23]
The first whole genome assembly was estimated to contain around 57,000 genes,[20] though the more recent genome sequences support estimates between 42,000 and 44,700 protein-coding genes.[22][23] The availability of whole genome sequences has provided evidence that the wild ancestor of the cultivated apple most likely is Malus sieversii. Re-sequencing of multiple accessions has supported this, while also suggesting extensive introgression from Malus sylvestris following domestication.[24]
Cultivation
History
Map of the origins of the cultivated apple. The wild origin is in Kazakhstan; hybridisations and repeated domestications followed, modifying many attributes of the fruit.[24]Wild Malus sieversii apple in Kazakhstan
Central Asia is generally considered the center of origin for apples due to the genetic variability in specimens there.[25] The wild ancestor of Malus domestica was Malus sieversii, found growing wild in the mountains of Central Asia in southern Kazakhstan, Kyrgyzstan, Tajikistan, and northwestern China.[5][26] Cultivation of the species, most likely beginning on the forested flanks of the Tian Shan mountains, progressed over a long period of time and permitted secondary introgression of genes from other species into the open-pollinated seeds. Significant exchange with Malus sylvestris, the crabapple, resulted in populations of apples being more related to crabapples than to the more morphologically similar progenitor Malus sieversii. In strains without recent admixture the contribution of the latter predominates.[27][28][29]
The apple is thought to have been domesticated 4,000–10,000 years ago in the Tian Shan mountains, and then to have travelled along the Silk Road to Europe, with hybridization and introgression of wild crabapples from Siberia (M. baccata), the Caucasus (M. orientalis), and Europe (M. sylvestris). Only the M. sieversii trees growing on the western side of the Tian Shan mountains contributed genetically to the domesticated apple, not the isolated population on the eastern side.[24]
Chinese soft apples, such as M. asiatica and M. prunifolia, have been cultivated as dessert apples for more than 2,000 years in China. These are thought to be hybrids between M. baccata and M. sieversii in Kazakhstan.[24]
Among the traits selected for by human growers are size, fruit acidity, color, firmness, and soluble sugar. Unusually for domesticated fruits, the wild M. sieversii origin is only slightly smaller than the modern domesticated apple.[24]
At the Sammardenchia-Cueis site near Udine in Northeastern Italy, seeds from some form of apples have been found in material carbon dated to between 6570 and 5684 BCE.[30] Genetic analysis has not yet been successfully used to determine whether such ancient apples were wild Malus sylvestris or Malus domesticus containing Malus sieversii ancestry. It is hard to distinguish in the archeological record between foraged wild apples and apple plantations.[31]
There is indirect evidence of apple cultivation in the third millennium BCE in the Middle East.[31] There is direct evidence, apple cores, dated to the 10th century BCE from a Judean site between the Sinai and Negev. [32] There was substantial apple production in European classical antiquity, and grafting was certainly known then.[31] Grafting is an essential part of modern domesticated apple production, to be able to propagate the best cultivars; it is unclear when apple tree grafting was invented.[31]
The Roman writer Pliny the Elder describes a method of storage for apples from his time in the 1st century. He says they should be placed in a room with good air circulation from a north facing window on a bed of straw, chaff, or mats with windfalls kept separately.[33] Though methods like this will extend the availabity of reasonably fresh apples, without refrigeration their lifespan is limited. Even sturdy winter apple varieties will only keep well until December in cool climates.[34] For longer storage medieval Europeans strung up cored and peeled apples to dry, either whole or sliced into rings.[35]
Of the many Old World plants that the Spanish introduced to Chiloé Archipelago in the 16th century, apple trees became particularly well adapted.[36] Apples were introduced to North America by colonists in the 17th century,[5] and the first named apple cultivar was introduced in Boston by Reverend William Blaxton in 1640.[37] The only apples native to North America are crab apples.[38]
Apple cultivars brought as seed from Europe were spread along Native American trade routes, as well as being cultivated on colonial farms. An 1845 United States apples nursery catalogue sold 350 of the “best” cultivars, showing the proliferation of new North American cultivars by the early 19th century.[38] In the 20th century, irrigation projects in Eastern Washington began and allowed the development of the multibillion-dollar fruit industry, of which the apple is the leading product.[5]
Until the 20th century, farmers stored apples in frostproof cellars during the winter for their own use or for sale. Improved transportation of fresh apples by train and road replaced the necessity for storage.[39][40]Controlled atmosphere facilities are used to keep apples fresh year-round. Controlled atmosphere facilities use high humidity, low oxygen, and controlled carbon dioxide levels to maintain fruit freshness. They were first researched at Cambridge University in the 1920s and first used in the United States in the 1950s.[41]
Many apples grow readily from seeds. However, apples must be propagated asexually to obtain cuttings with the characteristics of the parent. This is because seedling apples are “extreme heterozygotes“. Rather than resembling their parents, seedlings are all different from each other and from their parents.[42]Triploid cultivars have an additional reproductive barrier in that three sets of chromosomes cannot be divided evenly during meiosis, yielding unequal segregation of the chromosomes (aneuploids). Even in the case when a triploid plant can produce a seed (apples are an example), it occurs infrequently, and seedlings rarely survive.[43]
Because apples are not true breeders when planted as seeds, propagation usually involves grafting of cuttings. The rootstock used for the bottom of the graft can be selected to produce trees of a large variety of sizes, as well as changing the winter hardiness, insect and disease resistance, and soil preference of the resulting tree. Dwarf rootstocks can be used to produce very small trees (less than 3.0 m or 10 ft high at maturity), which bear fruit many years earlier in their life cycle than full size trees, and are easier to harvest.[44]
Dwarf rootstocks for apple trees can be traced as far back as 300 BCE, to the area of Persia and Asia Minor. Alexander the Great sent samples of dwarf apple trees to Aristotle‘s Lyceum. Dwarf rootstocks became common by the 15th century and later went through several cycles of popularity and decline throughout the world.[45] The majority of the rootstocks used to control size in apples were developed in England in the early 1900s. The East Malling Research Station conducted extensive research into rootstocks, and their rootstocks are given an “M” prefix to designate their origin. Rootstocks marked with an “MM” prefix are Malling-series cultivars later crossed with trees of ‘Northern Spy‘ in Merton, England.[46]
Most new apple cultivars originate as seedlings, which either arise by chance or are bred by deliberately crossing cultivars with promising characteristics.[47] The words “seedling”, “pippin”, and “kernel” in the name of an apple cultivar suggest that it originated as a seedling. Apples can also form bud sports (mutations on a single branch). Some bud sports turn out to be improved strains of the parent cultivar. Some differ sufficiently from the parent tree to be considered new cultivars.[48]
Apples have been acclimatized in Ecuador at very high altitudes, where they can often, with the needed factors, provide crops twice per year because of constant temperate conditions year-round.[49]
Apples are self-incompatible; they must cross-pollinate to develop fruit. During the flowering each season, apple growers often utilize pollinators to carry pollen. Honey bees are most commonly used. Orchard mason bees are also used as supplemental pollinators in commercial orchards. Bumblebeequeens are sometimes present in orchards, but not usually in sufficient number to be significant pollinators.[48][50]
Cultivars are sometimes classified by the day of peak bloom in the average 30-day blossom period, with pollinizers selected from cultivars within a 6-day overlap period. There are four to seven pollination groups in apples, depending on climate:
Group A – Early flowering, 1 to 3 May in England (‘Gravenstein‘, ‘Red Astrachan’)
Cultivars vary in their yield and the ultimate size of the tree, even when grown on the same rootstock. Some cultivars, if left unpruned, grow very large—letting them bear more fruit, but making harvesting more difficult. Depending on tree density (number of trees planted per unit surface area), mature trees typically bear 40–200 kg (90–440 lb) of apples each year, though productivity can be close to zero in poor years. Apples are harvested using three-point ladders that are designed to fit amongst the branches. Trees grafted on dwarfing rootstocks bear about 10–80 kg (20–180 lb) of fruit per year.[48]
Some farms with apple orchards open them to the public so consumers can pick their own apples.[52]
Crops ripen at different times of the year according to the cultivar. Cultivar that yield their crop in the summer include ‘Sweet Bough‘ and ‘Duchess’; fall producers include ‘Blenheim’; winter producers include ‘King’, ‘Swayzie‘, and ‘Tolman Sweet’.[38]
Commercially, apples can be stored for months in controlled atmosphere chambers. Apples are commonly stored in chambers with lowered concentrations of oxygen to reduce respiration and slow softening and other changes if the fruit is already fully ripe. The gas ethylene is used by plants as a hormone which promotes ripening, decreasing the time an apple can be stored. For storage longer than about six months the apples are picked earlier, before full ripeness, when ethylene production by the fruit is low. However, in many varieties this increases their sensitivity to carbon dioxide, which also must be controlled.[53]
For home storage, most culitvars of apple can be stored for three weeks in a pantry and four to six weeks from the date of purchase in a refrigerator that maintains 4 to 0 °C (39 to 32 °F).[54][55] Some varieties of apples (e.g. ‘Granny Smith‘ and ‘Fuji‘) have more than three times the storage life of others.[56]
Non-organic apples may be sprayed with a substance 1-methylcyclopropene blocking the apples’ ethylene receptors, temporarily preventing them from ripening.[57]
Apple trees are susceptible to fungal and bacterial diseases, and to damage by insect pests. Many commercial orchards pursue a program of chemical sprays to maintain high fruit quality, tree health, and high yields. These prohibit the use of synthetic pesticides, though some older pesticides are allowed. Organic methods include, for instance, introducing its natural predator to reduce the population of a particular pest.
A wide range of pests and diseases can affect the plant. Three of the more common diseases or pests are mildew, aphids, and apple scab.
Mildew is characterized by light grey powdery patches appearing on the leaves, shoots and flowers, normally in spring. The flowers turn a creamy yellow color and do not develop correctly. This can be treated similarly to Botrytis—eliminating the conditions that caused the disease and burning the infected plants are among recommended actions.[58]
Aphids are small insects with sucking mouthparts. Five species of aphids commonly attack apples: apple grain aphid, rosy apple aphid, apple aphid, spirea aphid, and the woolly apple aphid. The aphid species can be identified by color, time of year, and by differences in the cornicles (small paired projections from their rear).[59] Aphids feed on foliage using needle-like mouth parts to suck out plant juices. When present in high numbers, certain species reduce tree growth and vigor.[60]
Apple scab: Apple scab causes leaves to develop olive-brown spots with a velvety texture that later turn brown and become cork-like in texture. The disease also affects the fruit, which also develops similar brown spots with velvety or cork-like textures. Apple scab is spread through fungus growing in old apple leaves on the ground and spreads during warm spring weather to infect the new year’s growth.[61]
Among the most serious disease problems is a bacterial disease called fireblight, and three fungal diseases: Gymnosporangium rust, black spot,[62] and bitter rot.[63]Codling moths, and the apple maggots of fruit flies, cause serious damage to apple fruits, making them unsaleable. Young apple trees are also prone to mammal pests like mice and deer, which feed on the soft bark of the trees, especially in winter.[61] The larvae of the apple clearwing moth (red-belted clearwing) burrow through the bark and into the phloem of apple trees, potentially causing significant damage.[64]
There are more than 7,500 known cultivars (cultivated varieties) of apples.[65] Cultivars vary in their yield and the ultimate size of the tree, even when grown on the same rootstock.[66] Different cultivars are available for temperate and subtropical climates. The UK’s National Fruit Collection, which is the responsibility of the Department of Environment, Food, and Rural Affairs, includes a collection of over 2,000 cultivars of apple tree in Kent.[67] The University of Reading, which is responsible for developing the UK national collection database, provides access to search the national collection. The University of Reading’s work is part of the European Cooperative Programme for Plant Genetic Resources of which there are 38 countries participating in the Malus/Pyrus work group.[68]
The UK’s national fruit collection database contains much information on the characteristics and origin of many apples, including alternative names for what is essentially the same “genetic” apple cultivar. Most of these cultivars are bred for eating fresh (dessert apples), though some are cultivated specifically for cooking (cooking apples) or producing cider. Cider apples are typically too tart and astringent to eat fresh, but they give the beverage a rich flavor that dessert apples cannot.[69]
In Europe, apple breeding programs are conducted at places such as Julius Kühn-Institut, the German federal research center for cultivated plants.[70]
In the United States there are many apple breeding programs associated with universities. For instance, in the East, Cornell University has had a program operating since 1880 in Geneva, New York, while in the West, Washington State University started a program to support their home state’s apple industry in 1994.[71] Released by the University of Minnesota in 1991, the ‘Honeycrisp‘ has become famous for its crispness and juiciness, thereby commanding high market prices.[71] Unusually for a popular cultivar, the ‘Honeycrisp’ is not directly related to another popular apple cultivar but instead to two unsuccessful cultivars.[72] However, it is also difficult to grow and to store, prompting the industry to seek hybrids that not only appeal to consumers are also less costly for farmers to cultivate and last longer in storage.[73] By the 2020s, about half of the new apple varieties entering the market in the United States and Canada are ‘Honeycrisp’ progeny.[74] Such hybrids include the ‘SweeTango‘ (a cross between the ‘Honeycrisp’ and the ‘Zestar‘) introduced by the University of Minnesota in 2008 and the ‘Cosmic Crisp‘ (the ‘Honeycrisp’ and the ‘Enterprise‘) released by Washington State University in 2017.[73]
Less common apple cultivars from an orchard in Italy
Commercially popular apple cultivars are soft but crisp. Other desirable qualities in modern commercial apple breeding are a colorful skin, absence of russeting, ease of shipping, lengthy storage ability, high yields, disease resistance, common apple shape, and developed flavor.[66] Modern apples are generally sweeter than older cultivars, as popular tastes in apples have varied over time.[75] Most North Americans and Europeans favor crunchy, sweet, and subacid apples.[76] Nevertheless, tart apples maintain a strong minority following.[77] In the United States today, the most popular apple varieties are the ‘Ambrosia‘, ‘Honeycrisp’, and ‘Jazz‘, according Nielsen data,[73] while in Canada, the ‘Honeyscrisp’, ‘Ambrosia’, and ‘Gala’ take the top spots.[75] Together, these relatively newer varieties have overtaken once dominant cultivars like the ‘McIntosh‘ and the ‘Red Delicious‘ in the North American market.[75][76] Extremely sweet apples with barely any acid flavor are popular in Asia,[77] especially the Indian subcontinent.[69]
Old cultivars are often oddly shaped, russeted, and grow in a variety of textures and colors. Some find them to have better flavor than modern cultivars, but they may have other problems that make them commercially unviable—low yield, disease susceptibility, poor tolerance for storage or transport, or just being the “wrong” size.[78] A few old cultivars are still produced on a large scale, but many have been preserved by home gardeners and farmers that sell directly to local markets. Many unusual and locally important cultivars with their own unique taste and appearance exist; apple conservation campaigns have sprung up around the world to preserve such local cultivars from extinction. In the United Kingdom, old cultivars such as ‘Cox’s Orange Pippin‘ and ‘Egremont Russet‘ are still commercially important even though by modern standards they are low yielding and susceptible to disease.[5]
World production of apples in 2023 was 97 million tonnes, with China producing 51% of the total (table).[79] Secondary producers were the United States, Turkey, and Poland.[79]
Toxicity
Amygdalin
Apple seeds contain small amounts of amygdalin, a sugar and cyanide compound known as a cyanogenic glycoside. Ingesting small amounts of apple seeds causes no ill effects, but consumption of extremely large doses can cause adverse reactions. It may take several hours before the poison takes effect, as cyanogenic glycosides must be hydrolyzed before the cyanide ion is released.[80] The U.S. National Library of Medicine‘s Hazardous Substances Data Bank records no cases of amygdalin poisoning from consuming apple seeds.[81]
Allergy
One form of apple allergy, often found in northern Europe, is called birch-apple syndrome and is found in people who are also allergic to birchpollen.[82] Allergic reactions are triggered by a protein in apples that is similar to birch pollen, and people affected by this protein can also develop allergies to other fruits, nuts, and vegetables. Reactions, which entail oral allergy syndrome (OAS), generally involve itching and inflammation of the mouth and throat,[82] but in rare cases can also include life-threatening anaphylaxis.[83] This reaction only occurs when raw fruit is consumed—the allergen is neutralized in the cooking process. The variety of apple, maturity and storage conditions can change the amount of allergen present in individual fruits. Long storage times can increase the amount of proteins that cause birch-apple syndrome.[82]
In other areas, such as the Mediterranean, some individuals have adverse reactions to apples because of their similarity to peaches.[82] This form of apple allergy also includes OAS, but often has more severe symptoms, such as vomiting, abdominal pain and urticaria, and can be life-threatening. Individuals with this form of allergy can also develop reactions to other fruits and nuts. Cooking does not break down the protein causing this particular reaction, so affected individuals cannot eat raw or cooked apples. Freshly harvested, over-ripe fruits tend to have the highest levels of the protein that causes this reaction.[82]
Breeding efforts have yet to produce a hypoallergenic fruit suitable for either of the two forms of apple allergy.[82]
A raw apple is 86% water and 14% carbohydrates, with negligible content of fat and protein (table). A reference serving of a raw apple with skin weighing 100 g (3.5 oz) provides 52 calories and a moderate content of dietary fiber (table). Otherwise, there is low content of micronutrients, with the Daily Values of all falling below 10% (table).
Machine for paring, coring, and slicing apples, from Henry B. Scammell’s 1897 handbook Cyclopedia of Valuable Receipts
Apples varieties can be grouped as cooking apples, eating apples, and cider apples, the last so astringent as to be “almost inedible”.[86] Apples are consumed as juice, raw in salads, baked in pies, cooked into sauces and apple butter, or baked.[87] They are sometimes used as an ingredient in savory foods, such as sausage and stuffing.[88]
Several techniques are used to preserve apples and apple products. Traditional methods include drying and making apple butter.[86] Juice and cider are produced commercially; cider is a significant industry in regions such as the West of England and Normandy.[86]
Apples are an important ingredient in many desserts, such as pies, crumbles, and cakes. When cooked, some apple cultivars easily form a puree known as apple sauce, which can be cooked down to form a preserve, apple butter. They are often baked or stewed, and are cooked in some meat dishes.[86]
Apples are milled or pressed to produce apple juice, which may be drunk unfiltered (called apple cider in North America), or filtered. Filtered juice is often concentrated and frozen, then reconstituted later and consumed. Apple juice can be fermented to make cider (called hard cider in North America), ciderkin, and vinegar.[8] Through distillation, various alcoholic beverages can be produced, such as applejack, Calvados, and apple brandy.[8][91]
Organic production
Organic apples are commonly produced in the United States.[92] Due to infestations by key insects and diseases, organic production is difficult in Europe.[93] The use of pesticides containing chemicals, such as sulfur, copper, microorganisms, viruses, clay powders, or plant extracts (pyrethrum, neem) has been approved by the EU Organic Standing Committee to improve organic yield and quality.[93] A light coating of kaolin, which forms a physical barrier to some pests, also may help prevent apple sun scalding.[48]
In Norse mythology, the goddess Iðunn is portrayed in the Prose Edda (written in the 13th century by Snorri Sturluson) as providing apples to the gods that give them eternal youthfulness. The English scholar H. R. Ellis Davidson links apples to religious practices in Germanic paganism, from which Norse paganism developed. She points out that buckets of apples were found in the Oseberg ship burial site in Norway, that fruit and nuts (Iðunn having been described as being transformed into a nut in Skáldskaparmál) have been found in the early graves of the Germanic peoples in England and elsewhere on the continent of Europe, which may have had a symbolic meaning, and that nuts are still a recognized symbol of fertility in southwest England.[101]
Davidson notes a connection between apples and the Vanir, a tribe of gods associated with fertility in Norse mythology, citing an instance of eleven “golden apples” being given to woo the beautiful Gerðr by Skírnir, who was acting as messenger for the major Vanir god Freyr in stanzas 19 and 20 of Skírnismál. Davidson also notes a further connection between fertility and apples in Norse mythology in chapter 2 of the Völsunga saga: when the major goddess Frigg sends King Rerir an apple after he prays to Odin for a child, Frigg’s messenger (in the guise of a crow) drops the apple in his lap as he sits atop a mound.[101] Rerir’s wife’s consumption of the apple results in a six-year pregnancy and the birth (by Caesarean section) of their son—the hero Völsung.[102]
Further, Davidson points out the “strange” phrase “Apples of Hel” used in an 11th-century poem by the skald Thorbiorn Brúnarson. She states this may imply that the apple was thought of by Brúnarson as the food of the dead. Further, Davidson notes that the potentially Germanic goddess Nehalennia is sometimes depicted with apples and that parallels exist in early Irish stories. Davidson asserts that while cultivation of the apple in Northern Europe extends back to at least the time of the Roman Empire and came to Europe from the Near East, the native varieties of apple trees growing in Northern Europe are small and bitter. Davidson concludes that in the figure of Iðunn “we must have a dim reflection of an old symbol: that of the guardian goddess of the life-giving fruit of the other world.”[101]
The Greek goddess of discord, Eris, became disgruntled after she was excluded from the wedding of Peleus and Thetis.[105] In retaliation, she tossed a golden apple inscribed Καλλίστη (Kallistē, “For the most beautiful one”), into the wedding party. Three goddesses claimed the apple: Hera, Athena, and Aphrodite. Paris of Troy was appointed to select the recipient. After being bribed by both Hera and Athena, Aphrodite tempted him with the most beautiful woman in the world, Helen of Sparta. He awarded the apple to Aphrodite, thus indirectly causing the Trojan War.[106][107]
The apple was thus considered, in ancient Greece, sacred to Aphrodite. To throw an apple at someone was to symbolically declare one’s love; and similarly, to catch it was to symbolically show one’s acceptance of that love. An epigram claiming authorship by Plato states:[108]
I throw the apple at you, and if you are willing to love me, take it and share your girlhood with me; but if your thoughts are what I pray they are not, even then take it, and consider how short-lived is beauty.
Atalanta, also of Greek mythology, raced all her suitors in an attempt to avoid marriage. She outran all but Hippomenes (also known as Melanion, a name possibly derived from melon, the Greek word for both “apple” and fruit in general),[104] who defeated her by cunning, not speed. Hippomenes knew that he could not win in a fair race, so he used three golden apples (gifts of Aphrodite, the goddess of love) to distract Atalanta. It took all three apples and all of his speed, but Hippomenes was finally successful, winning the race and Atalanta’s hand.[109][110]
Píngānguǒ (“Peace apples”) on sale in Beijing for Christmas Eve (2017)
In China, apples symbolise peace, since the sounds of the first element (“píng”) in the words “apple” (苹果, Píngguǒ) and “peace” (平安, Píng’ān) are homophonous in Mandarin and Cantonese.[3][112] When these two words are combined, the word Píngānguǒ (平安果, “Peace apples”) is formed. This association developed further as the name for Christmas Eve in Mandarin is Píngānyè (平安夜, “Peaceful/Quiet Evening”), which made the gifting of apples at this season to friends and associates popular, as a way to wish them peace and safety.[112]
Though the forbidden fruit of Eden in the Book of Genesis is not identified, popular Christian tradition has held that it was an apple that Eve coaxed Adam to share with her.[113] The origin of the popular identification with a fruit unknown in the Middle East in biblical times is found in wordplay with the Latin words mālum (an apple) and mălum (an evil), each of which is normally written malum.[114] The tree of the forbidden fruit is called “the tree of the knowledge of good and evil” in Genesis 2:17,[115] and the Latin for “good and evil” is bonum et malum.[116]
Renaissance painters may also have been influenced by the story of the golden apples in the Garden of Hesperides. As a result, in the story of Adam and Eve, the apple became a symbol for knowledge, immortality, temptation, the fall of man into sin, and sin itself. The larynx in the human throat has been called the “Adam’s apple” because of a notion that it was caused by the forbidden fruit remaining in the throat of Adam. The apple as symbol of sexual seduction has been used to imply human sexuality, possibly in an ironic vein.[113]
Proverb
The proverb, “An apple a day keeps the doctor away“, addressing the supposed health benefits of the fruit, has been traced to 19th-century Wales, where the original phrase was “Eat an apple on going to bed, and you’ll keep the doctor from earning his bread”.[117] In the 19th century and early 20th, the phrase evolved to “an apple a day, no doctor to pay” and “an apple a day sends the doctor away”; the phrasing now commonly used was first recorded in 1922.[118]
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