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An apple is an edible
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
produced by an apple tree (''Malus domestica''). Apple
trees In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are u ...
are cultivated worldwide and are the most widely grown species in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''
Malus ''Malus'' ( or ) is a genus of about 30–55 species of small deciduous trees or shrubs in the family Rosaceae, including the domesticated orchard apple, crab apples, wild apples, and rainberries. The genus is native to the temperate zone ...
''. The
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
originated in
Central Asia Central Asia, also known as Middle Asia, is a subregion, region of Asia that stretches from the Caspian Sea in the west to western China and Mongolia in the east, and from Afghanistan and Iran in the south to Russia in the north. It includes t ...
, where its wild ancestor, ''
Malus sieversii ''Malus sieversii'' is a wild apple native to the mountains of Central Asia in southern Kazakhstan. It has recently been shown to be the primary ancestor of most cultivars of the domesticated apple (''Malus domestica''). It was first described a ...
'', is still found today. Apples have been grown for thousands of years in Asia and Europe and were brought to North America by European colonists. Apples have
religious Religion is usually defined as a social- cultural system of designated behaviors and practices, morals, beliefs, worldviews, texts, sanctified places, prophecies, ethics, or organizations, that generally relates humanity to supernatur ...
and mythological significance in many cultures, including Norse,
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
, and
European Christian Christianity is the largest religion in Europe. Christianity has been practiced in Europe since the first century, and a number of the Pauline Epistles were addressed to Christians living in Greece, as well as other parts of the Roman Empire. ...
tradition. Apples grown from seed tend to be very different from those of their parents, and the resultant fruit frequently lacks desired characteristics. Generally, apple
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture ...
s are propagated by clonal grafting onto
rootstock A rootstock is part of a plant, often an underground part, from which new above-ground growth can be produced. It could also be described as a stem with a well developed root system, to which a bud from another plant is grafted. It can refer to a ...
s. 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 known cultivars of apples. Different cultivars are bred for various tastes and uses, including cooking, eating raw, and cider production. Trees and fruit are prone to a number of
fungal A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
, bacterial, and pest problems, which can be controlled by a number of organic and non-organic means. In 2010, the fruit's
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding g ...
was
sequenced In genetics and biochemistry, sequencing means to determine the primary structure (sometimes incorrectly called the primary sequence) of an unbranched biopolymer. Sequencing results in a symbolic linear depiction known as a sequence which suc ...
as part of research on disease control and selective breeding in apple production. Worldwide production of apples in 2018 was 86 million
tonnes The tonne ( or ; symbol: t) is a unit of mass equal to 1000 kilograms. It is a non-SI unit accepted for use with SI. It is also referred to as a metric ton to distinguish it from the non-metric units of the short ton (United States c ...
, with China accounting for nearly half of the total.


Etymology

The word ''apple'', whose
Old English Old English (, ), or Anglo-Saxon, is the earliest recorded form of the English language, spoken in England and southern and eastern Scotland in the early Middle Ages. It was brought to Great Britain by Anglo-Saxon settlers in the mid-5th c ...
ancestor is , is descended from the
Proto-Germanic Proto-Germanic (abbreviated PGmc; also called Common Germanic) is the reconstructed proto-language of the Germanic branch of the Indo-European languages. Proto-Germanic eventually developed from pre-Proto-Germanic into three Germanic bran ...
noun , descended in turn from
Proto-Indo-European Proto-Indo-European (PIE) is the reconstructed common ancestor of the Indo-European language family. Its proposed features have been derived by linguistic reconstruction from documented Indo-European languages. No direct record of Proto-Indo- ...
. As late as the 17th century, the word also functioned as a generic term for all fruit including
nut Nut often refers to: * Nut (fruit), fruit composed of a hard shell and a seed, or a collective noun for dry and edible fruits or seeds * Nut (hardware), fastener used with a bolt Nut or Nuts may also refer to: Arts, entertainment, and media Co ...
s—such as the 14th-century
Middle English Middle English (abbreviated to ME) is a form of the English language that was spoken after the Norman conquest of 1066, until the late 15th century. The English language underwent distinct variations and developments following the Old English ...
expression , meaning a banana. This use is analogous to the French language use of .


Description

The apple is a
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, ...
tree, generally standing tall in cultivation and up to in the wild. When cultivated, the size, shape and branch density are determined by
rootstock A rootstock is part of a plant, often an underground part, from which new above-ground growth can be produced. It could also be described as a stem with a well developed root system, to which a bud from another plant is grafted. It can refer to a ...
selection and trimming method. The leaves are alternately arranged dark green-colored simple ovals with serrated margins and slightly downy undersides.
Blossoms In botany, blossoms are the flowers of stone fruit trees (genus ''Prunus'') and of some other plants with a similar appearance that flower profusely for a period of time in spring. Colloquially, flowers of orange are referred to as such as wel ...
are produced in
spring Spring(s) may refer to: Common uses * Spring (season) Spring, also known as springtime, is one of the four temperate seasons, succeeding winter and preceding summer. There are various technical definitions of spring, but local usage of ...
simultaneously with the budding of the leaves and are produced on spurs and some long
shoot In botany, a plant shoot consists of any plant stem together with its appendages, leaves and lateral buds, flowering stems, and flower buds. The new growth from seed germination that grows upward is a shoot where leaves will develop. In the sp ...
s. The flowers are white with a pink tinge that gradually fades, five petaled, with an
inflorescence An inflorescence is a group or cluster of flowers arranged on a Plant stem, stem that is composed of a main branch or a complicated arrangement of branches. Morphology (biology), Morphologically, it is the modified part of the shoot of sperma ...
consisting of a cyme with 4–6 flowers. The central flower of the inflorescence is called the "king bloom"; it opens first and can develop a larger fruit. The
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
is a pome that matures in late summer or autumn, and cultivars exist in a wide range of sizes. Commercial growers aim to produce an apple that is in diameter, due to market preference. Some consumers, especially those in Japan, prefer a larger apple, while apples below are generally used for making juice and have little fresh market value. The skin of ripe apples is generally red, yellow, green, pink, or
russetted Russeting or russetting is an abnormality of fruit skin which manifests in russet-colored (brownish) patches that are rougher than healthy skin. It is a common feature in apples and pears. Russeting is typically an undesirable trait, which reduce ...
, though many bi- or tri-colored cultivars may be found. The skin may also be wholly or partly russeted i.e. rough and brown. The skin is covered in a protective layer of
epicuticular wax Epicuticular wax is a coating of wax covering the outer surface of the plant cuticle in land plants. It may form a whitish film or bloom on leaves, fruits and other plant organs. Chemically, it consists of hydrophobic organic compounds, mainly st ...
. The exocarp (flesh) is generally pale yellowish-white, though pink, yellow or green exocarps also occur.


Wild ancestors

The original wild ancestor of ''Malus domestica'' was ''Malus sieversii'', found growing wild in the
mountains of Central Asia The Mountains of Central Asia are a biodiversity hot spot designated by Conservation International which covers several montane and alpine ecoregions of Central Asia, including those of the Pamir and Tian Shan ranges, and extending across portio ...
in southern
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a transcontinental country located mainly in Central Asia and partly in Eastern Europe. It borders Russia to the north and west, China to the east, Kyrgyzstan to the southeast, Uzbeki ...
,
Kyrgyzstan Kyrgyzstan,, pronounced or the Kyrgyz Republic, is a landlocked country in Central Asia. Kyrgyzstan is bordered by Kazakhstan to the north, Uzbekistan to the west, Tajikistan to the south, and the People's Republic of China to the ea ...
,
Tajikistan Tajikistan (, ; tg, Тоҷикистон, Tojikiston; russian: Таджикистан, Tadzhikistan), officially the Republic of Tajikistan ( tg, Ҷумҳурии Тоҷикистон, Jumhurii Tojikiston), is a landlocked country in Centr ...
, and
northwestern China Northwest China () is a statistical region of China which includes the autonomous regions of Xinjiang and Ningxia and the provinces of Shaanxi, Gansu and Qinghai. It has an area of 3,107,900 km2. The region is characterized by a (semi-)arid con ...
. Cultivation of the species, most likely beginning on the forested flanks of the
Tian Shan The Tian Shan,, , otk, 𐰴𐰣 𐱅𐰭𐰼𐰃, , tr, Tanrı Dağı, mn, Тэнгэр уул, , ug, تەڭرىتاغ, , , kk, Тәңіртауы / Алатау, , , ky, Теңир-Тоо / Ала-Тоо, , , uz, Tyan-Shan / Tangritog‘ ...
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 ''Malus sylvestris'', the European crab apple, is a species of the genus ''Malus'', native to Europe. Its scientific name means "forest apple" and the truly wild tree has thorns. Description Wild apple has an expanded crown and often appears mo ...
'', the crabapple, resulted in current 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.


Genome

Apples are diploid (though triploid cultivars are not uncommon), have 17 chromosomes and an estimated
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding g ...
size of approximately 650 Mb. Several whole genome sequences have been made available, the first one in 2010 was based on the diploid cultivar ' Golden Delicious'. However, this first whole genome sequence turned out to contain several errors 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. The first whole genome assembly was estimated to contain around 57,000 genes, though the more recent genome sequences support more moderate estimates between 42,000 and 44,700 protein-coding genes. Among other things, 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.


History

''
Malus sieversii ''Malus sieversii'' is a wild apple native to the mountains of Central Asia in southern Kazakhstan. It has recently been shown to be the primary ancestor of most cultivars of the domesticated apple (''Malus domestica''). It was first described a ...
'' is recognized as a major progenitor species to the cultivated apple, and is morphologically similar. Due to the genetic variability in Central Asia, this region is generally considered the center of origin for apples. The apple is thought to have been domesticated 4000–10000 years ago in the
Tian Shan The Tian Shan,, , otk, 𐰴𐰣 𐱅𐰭𐰼𐰃, , tr, Tanrı Dağı, mn, Тэнгэр уул, , ug, تەڭرىتاغ, , , kk, Тәңіртауы / Алатау, , , ky, Теңир-Тоо / Ала-Тоо, , , uz, Tyan-Shan / Tangritog‘ ...
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. Chinese soft apples, such as '' M. asiatica'' and '' M. prunifolia'', have been cultivated as dessert apples for more than 2000 years in China. These are thought to be hybrids between ''M. baccata'' and ''M. sieversii'' in Kazakhstan. 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. At the Sammardenchia-Cueis site near Udine in Northeastern Italy, seeds from some form of apples have been found in material carbon dated to around 4000 BCE. 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 generally also hard to distinguish in the archeological record between foraged wild apples and apple plantations. There is indirect evidence of apple cultivation in the third millennium BCE in the
Middle East The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabia (including the Arabian Peninsula and Bahrain), Asia Minor (Asian part of Turkey except Hatay Province), East Thrace (Europ ...
. There was substantial apple production in the European classical antiquity, and grafting was certainly known then. 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. Winter apples, picked in late autumn and stored just above freezing, have been an important food in Asia and Europe for millennia. Of the many Old World plants that the Spanish introduced to
Chiloé Archipelago The Chiloé Archipelago ( es, Archipiélago de Chiloé, , ) is a group of islands lying off the coast of Chile, in the Los Lagos Region. It is separated from mainland Chile by the Chacao Channel in the north, the Sea of Chiloé in the east and t ...
in the 16th century, apple trees became particularly well adapted. Apples were introduced to North America by colonists in the 17th century, and the first apple orchard on the North American continent was planted in
Boston Boston (), officially the City of Boston, is the state capital and most populous city of the Commonwealth of Massachusetts, as well as the cultural and financial center of the New England region of the United States. It is the 24th- mo ...
by Reverend
William Blaxton Reverend William Blaxton (also spelled William Blackstone) (1595 – 26 May 1675) was an early English settler in New England and the first European settler of Boston and Rhode Island. Biography William Blaxton was born in Horncastle, Lincoln ...
in 1625. The only apples native to North America are crab apples, which were once called "common apples". 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. In the 20th century, irrigation projects in
Eastern Washington Eastern Washington is the region of the U.S. state of Washington located east of the Cascade Range. It contains the city of Spokane (the second largest city in the state), the Tri-Cities, the Columbia River and the Grand Coulee Dam, the Hanf ...
began and allowed the development of the multibillion-dollar fruit industry, of which the apple is the leading product. 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.
Controlled atmosphere A controlled atmosphere is an agricultural storage method in which the concentrations of oxygen, carbon dioxide and nitrogen, as well as the temperature and humidity of a storage room are regulated. Both dry commodities and fresh fruit and vegetab ...
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 used in the United States in the 1960s.


Significance in European cultures and societies


Germanic paganism

In Norse mythology, the goddess
Iðunn In Norse mythology, Iðunn is a goddess associated with apples and youth. Iðunn is attested in the ''Poetic Edda'', compiled in the 13th century from earlier traditional sources, and the ''Prose Edda'', written in the 13th century by Snorri St ...
is portrayed in the ''
Prose Edda The ''Prose Edda'', also known as the ''Younger Edda'', ''Snorri's Edda'' ( is, Snorra Edda) or, historically, simply as ''Edda'', is an Old Norse textbook written in Iceland during the early 13th century. The work is often assumed to have been t ...
'' (written in the 13th century by Snorri Sturluson) as providing apples to the
gods A deity or god is a supernatural being who is considered divine or sacred. The ''Oxford Dictionary of English'' defines deity as a god or goddess, or anything revered as divine. C. Scott Littleton defines a deity as "a being with powers greater ...
that give them
eternal youth Eternal youth is the concept of human physical immortality free of ageing. The youth referred to is usually meant to be in contrast to the depredations of aging, rather than a specific age of the human lifespan. Eternal youth is common in myth ...
fulness. The English scholar H. R. Ellis Davidson links apples to religious practices in
Germanic paganism Germanic paganism or Germanic religion refers to the traditional, culturally significant religion of the Germanic peoples. With a chronological range of at least one thousand years in an area covering Scandinavia, the British Isles, modern Germ ...
, from which
Norse paganism Old Norse religion, also known as Norse paganism, is the most common name for a branch of Germanic religion which developed during the Proto-Norse period, when the North Germanic peoples separated into a distinct branch of the Germanic peop ...
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 ''Skáldskaparmál'' (Old Norse: 'The Language of Poetry'; c. 50,000 words; ; ) is the second part of the ''Prose Edda''. The section consists of a dialogue between Ægir, the divine personification of the sea, and Bragi, the god of poetry, ...
'') have been found in the early graves of the
Germanic peoples The Germanic peoples were historical groups of people that once occupied Central Europe and Scandinavia during antiquity and into the early Middle Ages. Since the 19th century, they have traditionally been defined by the use of ancient and e ...
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 Fertility is the capability to produce offspring through reproduction following the onset of sexual maturity. The fertility rate is the average number of children born by a female during her lifetime and is quantified demographically. Fertili ...
in southwest England. Davidson notes a connection between apples and the
Vanir In Norse mythology, the Vanir (; Old Norse: , singular Vanr ) are a group of gods associated with fertility, wisdom, and the ability to see the future. The Vanir are one of two groups of gods (the other being the Æsir) and are the namesake of the ...
, a tribe of gods associated with
fertility Fertility is the capability to produce offspring through reproduction following the onset of sexual maturity. The fertility rate is the average number of children born by a female during her lifetime and is quantified demographically. Fertili ...
in Norse mythology, citing an instance of eleven "golden apples" being given to woo the beautiful
Gerðr In Norse mythology, Gerðr (Old Norse: ; "fenced-in"Orchard (1997:54).) is a jötunn, goddess, and the wife of the god Freyr. Gerðr is attested in the ''Poetic Edda'', compiled in the 13th century from earlier traditional sources; the ''Prose Ed ...
by
Skírnir In Norse mythology, Skírnir (Old Norse" ; "bright one"Orchard (1997:149).) is the god Freyr's messenger and vassal. In the ''Poetic Edda'' poem ''Skírnismál'', Skírnir is sent as a messenger to Jötunheimr to conduct lovesick Freyr's wooing o ...
, who was acting as messenger for the major Vanir god
Freyr Freyr (Old Norse: 'Lord'), sometimes anglicized as Frey, is a widely attested god in Norse mythology, associated with kingship, fertility, peace, and weather. Freyr, sometimes referred to as Yngvi-Freyr, was especially associated with Sweden an ...
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 The ''Völsunga saga'' (often referred to in English as the ''Volsunga Saga'' or ''Saga of the Völsungs'') is a legendary saga, a late 13th-century poetic rendition in Old Norse of the origin and decline of the Völsung clan (including the st ...
'': when the major goddess
Frigg Frigg (; Old Norse: ) is a goddess, one of the Æsir, in Germanic mythology. In Norse mythology, the source of most surviving information about her, she is associated with marriage, prophecy, clairvoyance and motherhood, and dwells in the wet ...
sends King
Rerir In ''Völsunga saga'', Rerir, the son of Sigi, succeeds his murdered father and avenges his death. He rules in Hunaland and becomes a powerful ruler. Rerir's son is Völsung. Rerir and his wife were unable to have children until the goddess Fri ...
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 A mound is a heaped pile of earth, gravel, sand, rocks, or debris. Most commonly, mounds are earthen formations such as hills and mountains, particularly if they appear artificial. A mound may be any rounded area of topographically higher ...
. 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 In Norse mythology, Völsung ( non, Vǫlsungr ) was the son of Rerir and the eponymous ancestor of the ill-fated Völsung clan (), which includes the well known Norse hero Sigurð. He was murdered by the Geatish king Siggeir and later avenged b ...
. 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 Nehalennia (spelled variously) is a goddess of unclear origin, perhaps Germanic or Celtic. She is attested on and depicted upon numerous votive altars discovered around what is now the province of Zeeland, the Netherlands, where the Schelde Riv ...
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 The Roman Empire ( la, Imperium Romanum ; grc-gre, Βασιλεία τῶν Ῥωμαίων, Basileía tôn Rhōmaíōn) was the post- Republican period of ancient Rome. As a polity, it included large territorial holdings around the Mediter ...
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."


Greek mythology

Apples appear in many
religious traditions Religion is usually defined as a social-cultural system of designated behaviors and practices, morals, beliefs, worldviews, texts, sanctified places, prophecies, ethics, or organizations, that generally relates humanity to supernatural, tra ...
, often as a mystical or
forbidden fruit Forbidden fruit is a name given to the fruit growing in the Garden of Eden which God commands mankind not to eat. In the biblical story, Adam and Eve eat the fruit from the tree of the knowledge of good and evil and are exiled from Eden. As a ...
. One of the problems identifying apples in religion,
mythology Myth is a folklore genre consisting of narratives that play a fundamental role in a society, such as foundational tales or origin myths. Since "myth" is widely used to imply that a story is not objectively true, the identification of a narra ...
and folktales is that the word "apple" was used as a generic term for all (foreign) fruit, other than berries, including nuts, as late as the 17th century. For instance, in
Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the origin and nature of the world, the lives and activities ...
, the
Greek hero Hero cults were one of the most distinctive features of ancient Greek religion. In Homeric Greek, "hero" (, ) refers to the mortal offspring of a human and a god. By the historical period, however, the word came to mean specifically a ''dead'' ma ...
Heracles Heracles ( ; grc-gre, Ἡρακλῆς, , glory/fame of Hera), born Alcaeus (, ''Alkaios'') or Alcides (, ''Alkeidēs''), was a divine hero in Greek mythology, the son of Zeus and Alcmene, and the foster son of Amphitryon.By his adoptiv ...
, as a part of his
Twelve Labours The Labours of Hercules or Labours of Heracles ( grc-gre, wikt:ὁ, οἱ wikt:Ἡρακλῆς, Ἡρακλέους wikt:ἆθλος, ἆθλοι, ) are a series of episodes concerning a penance carried out by Heracles, the greatest of the ...
, was required to travel to the Garden of the Hesperides and pick the golden apples off the
Tree of Life The tree of life is a fundamental archetype in many of the world's mythological, religious, and philosophical traditions. It is closely related to the concept of the sacred tree.Giovino, Mariana (2007). ''The Assyrian Sacred Tree: A Hist ...
growing at its center. The Greek goddess of discord, Eris, became disgruntled after she was excluded from the wedding of
Peleus In Greek mythology, Peleus (; Ancient Greek: Πηλεύς ''Pēleus'') was a hero, king of Phthia, husband of Thetis and the father of their son Achilles. This myth was already known to the hearers of Homer in the late 8th century BC. Biogra ...
and
Thetis Thetis (; grc-gre, Θέτις ), is a figure from Greek mythology with varying mythological roles. She mainly appears as a sea nymph, a goddess of water, or one of the 50 Nereids, daughters of the ancient sea god Nereus. When described as ...
. In retaliation, she tossed a
golden apple The golden apple is an element that appears in various national and ethnic folk legends or fairy tales. Recurring themes depict a hero (for example Hercules or Făt-Frumos) retrieving the golden apples hidden or stolen by a monstrous antagonist. ...
inscribed Καλλίστη (''Kalliste'', sometimes transliterated ''Kallisti'', "For the most beautiful one"), into the wedding party. Three goddesses claimed the apple: Hera,
Athena Athena or Athene, often given the epithet Pallas, is an ancient Greek religion, ancient Greek goddess associated with wisdom, warfare, and handicraft who was later syncretism, syncretized with the Roman goddess Minerva. Athena was regarded ...
, and
Aphrodite Aphrodite ( ; grc-gre, Ἀφροδίτη, Aphrodítē; , , ) is an ancient Greek goddess associated with love, lust, beauty, pleasure, passion, and procreation. She was syncretized with the Roman goddess . Aphrodite's major symbols inclu ...
.
Paris Paris () is the Capital city, capital and List of communes in France with over 20,000 inhabitants, most populous city of France, with an estimated population of 2,165,423 residents in 2019 in an area of more than 105 km² (41 sq mi), ma ...
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 Troy, Helen of Sparta. He awarded the apple to Aphrodite, thus indirectly causing the Trojan War. 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: 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), 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.


Christian art

Though the forbidden fruit of Garden of Eden, 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. The origin of the popular identification with a fruit unknown in the Middle East in biblical times is found in confusion between the Latin language, Latin words ''mālum'' (an apple) and ''mălum'' (an evil), each of which is normally written ''malum''. The tree of the forbidden fruit is called "the tree of the knowledge of good and evil" in Genesis 2:17, and the Latin for "good and evil" is ''bonum et malum''. Renaissance painters may also have been influenced by the story of the golden apples in the Hesperides#The Garden of the Hesperides, 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.


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". 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.


Cultivars

There are more than 7,500 known
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture ...
s (cultivated varieties) of apples. Cultivars vary in their Crop yield, yield and the ultimate size of the tree, even when grown on the same
rootstock A rootstock is part of a plant, often an underground part, from which new above-ground growth can be produced. It could also be described as a stem with a well developed root system, to which a bud from another plant is grafted. It can refer to a ...
. 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. 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. 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. Commercially popular apple cultivars are soft but crisp. Other desirable qualities in modern commercial apple breeding are a colorful skin, absence of Russet apple, russeting, ease of shipping, lengthy storage ability, high yields, disease resistance, common apple shape, and developed flavor. Modern apples are generally sweeter than older cultivars, as popular tastes in apples have varied over time. Most North Americans and Europeans favor sweet, subacid apples, but tart apples have a strong minority following. Extremely sweet apples with barely any acid flavor are popular in Asia, especially the Indian subcontinent. 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. 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. File:Alice (apple).jpg, 'Alice' File:Ambrosia apples 2017 A3.jpg, 'Ambrosia (apple), Ambrosia' File:Malus-Ananasrenette.jpg, 'Ananasrenette' File:Arkansas Black apples (cropped).jpg, 'Arkansas Black' File:Aroma (apple).jpg, 'Aroma' File:Malus-Boskoop organic.jpg, 'Belle de Boskoop' File:Bramley's Seedling Apples.jpg, 'Bramley' File:Cox orange renette2.JPG, 'Cox's Orange Pippin' File:Cox’s Pomona.jpg, 'Cox Pomona' File:Pink Lady Apple (4107712628).jpg, 'Cripps Pink' File:Discovery apples.jpg, 'Discovery (apple), Discovery' File:Cross section of Egremont Russet (EMLA 1), National Fruit Collection (acc. 1979-159).jpg, 'Egremont Russet' File:Fuji apple.jpg, 'Fuji (apple), Fuji' File:Red Apple.jpg, 'Gala (apple), Gala' File:Jabuke Gloster na stablu.2c.jpg, 'Gloster (apple), Gloster' File:Golden Delicious apple 2021 (2).jpg, ' Golden Delicious' File:Apfel-Berlepsch.jpg, 'Goldrenette' (Reinette) File:Granny smith closeup.jpg, 'Granny Smith' File:Honeycrisp.jpg, 'Honeycrisp' File:Malus-James-Grieve.jpg, 'James Grieve' File:Jonagold.jpg, 'Jonagold' File:Malus Lobo 4397.jpg, 'Lobo' File:McIntosh.jpg, 'McIntosh (apple), McIntosh' File:Apple 03.jpg, 'Sciros' File:Red Delicious.jpg, 'Red Delicious' File:Shampion 13.01.2013 13-17-30.jpg, 'Champion (apple), Sampion' (Shampion) File:Mele stark.jpg, 'Stark Delicious' File:The SugarBee Apple now grown in Washington State.jpg, 'SugarBee' File:Summerred.jpg, 'Summerred' File:Tellissaare.JPG, 'Tellissaare' File:Yellow Transparent.jpg, 'Yellow Transparent'


Cultivation


Breeding

Many apples grow readily from seeds. However, more than with most perennial fruits, apples must be propagated asexually to obtain the sweetness and other desirable characteristics of the parent. This is because seedling apples are an example of "Zygosity, extreme heterozygotes", in that rather than inheriting genes from their parents to create a new apple with parental characteristics, they are instead significantly different from their parents, perhaps to compete with the many pests. Polyploid, 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. Because apples are not True-breeding organism, true breeders when planted as seeds, although cutting (plant), cuttings can take root and breed true, and may live for a century, grafting is usually used. The
rootstock A rootstock is part of a plant, often an underground part, from which new above-ground growth can be produced. It could also be described as a stem with a well developed root system, to which a bud from another plant is grafted. It can refer to a ...
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 high at maturity), which bear fruit many years earlier in their life cycle than full size trees, and are easier to harvest. 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. The majority of the rootstocks used today to control size in apples were developed in England in the early 1900s. The East Malling Research Station conducted extensive research into rootstocks, and today 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 London Borough of Merton, Merton, England. Most new apple cultivars originate as seedlings, which either arise by chance or are bred by deliberately crossing cultivars with promising characteristics. 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. Since the 1930s, the Excelsior Experiment Station at the University of Minnesota has introduced a steady progression of important apples that are widely grown, both commercially and by local orchardists, throughout Minnesota and Wisconsin. Its most important contributions have included 'Haralson (apple), Haralson' (which is the most widely cultivated apple in Minnesota), 'Wealthy (apple), Wealthy', 'Honeygold', and 'Honeycrisp'. 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.


Pollination

Apples are self-incompatible; they must Pollination, cross-pollinate to develop fruit. During the flowering each season, apple growers often utilize pollinators to carry pollen. Honey bees are most commonly used. Osmia lignaria, Orchard mason bees are also used as supplemental pollinators in commercial orchards. Bumblebee queen bee, queens are sometimes present in orchards, but not usually in sufficient number to be significant pollinators.Adamson, Nancy Lee
An Assessment of Non-Apis Bees as Fruit and Vegetable Crop Pollinators in Southwest Virginia
. Diss. 2011. Web. 15 October 2015.
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') * Group B – 4 to 7 May ('Idared', 'McIntosh (apple), McIntosh') * Group C – Mid-season flowering, 8 to 11 May ('Granny Smith', 'Cox's Orange Pippin') * Group D – Mid/late season flowering, 12 to 15 May (' Golden Delicious', 'Calville blanc d'hiver') * Group E – Late flowering, 16 to 18 May ('Braeburn', 'Reinette d'Orléans') * Group F – 19 to 23 May ('Suntan') * Group H – 24 to 28 May ('Court-Pendu Gris' – also called Court-Pendu plat) One cultivar can be pollinated by a compatible cultivar from the same group or close (A with A, or A with B, but not A with C or D). Cultivars are sometimes classified by the day of peak bloom in the average 30-day blossom period, with pollenizers selected from cultivars within a 6-day overlap period.


Maturation and harvest

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 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 of fruit per year. Farms with apple orchards open them to the public so consumers can pick their own apples. Crops ripen at different times of the year according to the cultivar. Cultivar that yield their crop in the summer include 'Gala', 'Golden Supreme', 'McIntosh', 'Transparent', 'Primate', 'Sweet Bough', and 'Duchess'; fall producers include 'Fuji', 'Jonagold', 'Golden Delicious', 'Red Delicious', 'Chenango', 'Gravenstein', 'Wealthy', 'McIntosh', 'Snow', and 'Blenheim'; winter producers include 'Winesap', 'Granny Smith', 'King', 'Wagener', 'Swazie (apple), Swayzie', 'Greening', and 'Tolman Sweet'.


Storage

Commercially, apples can be stored for a few months in controlled atmosphere chambers to delay ethylene-induced ripening. Apples are commonly stored in chambers with higher concentrations of carbon dioxide and high air filtration. This prevents ethylene concentrations from rising to higher amounts and preventing ripening from occurring too quickly. For home storage, most cultivars of apple can be held for approximately two weeks when kept at the coolest part of the refrigerator (i.e. below 5 °C). Some can be stored up to a year without significant degradation. Some varieties of apples (e.g. 'Granny Smith' and 'Fuji (apple), Fuji') have more than three times the storage life of others. Non-organic apples may be sprayed with a substance 1-methylcyclopropene blocking the apples' ethylene receptors, temporarily preventing them from ripening.


Pests and diseases

Apple trees are susceptible to a number of
fungal A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
and bacterium, bacterial diseases and 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 farming, 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 Botryotinia, ''Botrytis''—eliminating the conditions that caused the disease and burning the infected plants are among recommended actions. * Aphids are a small insect. 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). 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. * 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. Among the most serious disease problems is a bacterial disease called fireblight, and two fungal diseases: ''Gymnosporangium'' rust and Black Spot (disease), black spot. Other pests that affect apple trees include Codling moths and apple maggots. 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. The larvae of the Synanthedon myopaeformis, apple clearwing moth (red-belted clearwing) burrow through the bark and into the phloem of apple trees, potentially causing significant damage.


Production

World production of apples in 2020 was 86 million tonnes, with China producing 47% of the total (table). Secondary producers were the United States and Turkey.


Nutrition

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 grams provides 52 calories and a moderate content of dietary fiber. Otherwise, there is low content of micronutrients, with the Daily Values of all falling below 10%.


Uses

All parts of the fruit, including the skin, except for the seeds, are suitable for human consumption. The core, from stem to bottom, containing the seeds, is usually not eaten and is discarded. Apples can be consumed in various ways: Apple juice, juice, raw in salads, baked in Apple pie, pies, cooked into Apple sauce, sauces and spreads like apple butter, and other Cooking apple, baked dishes. Apples are sometimes used as an ingredient in savory foods, such as sausage and stuffing. Several techniques are used to preserve apples and apple products. Apples can be canned, dried or frozen. Canned or frozen apples are eventually baked into pies or other cooked dishes. Apple juice or cider is also bottled. Apple juice is often concentrated and frozen.


Popular uses

Apples are often eaten raw. Cultivars bred for raw consumption are termed dessert or table apples. * In the UK, a toffee apple is a traditional confection made by coating an apple in hot toffee and allowing it to cool. Similar treats in the U.S. are candy apples (coated in a hard shell of crystallized sugar syrup) and caramel apples (coated with cooled caramel). * Apples are eaten with honey at the Jewish New Year of Rosh Hashanah to symbolize a sweet new year. Apples are an important ingredient in many desserts, such as apple pie, apple crumble, apple crisp and apple cake. When cooked, some apple cultivars easily form a puree known as apple sauce. Apples are also made into apple butter and apple jelly. They are often baked or stewed and are also (cooked) in some meat dishes. Dried apples can be eaten or reconstituted (soaked in water, alcohol or some other liquid). 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 fermentation (food), fermented to make cider (called hard cider in North America), ciderkin, and vinegar. Through distillation, various alcoholic beverages can be produced, such as applejack (beverage), applejack, Calvados (spirit), Calvados, and apfelwein.


Organic production

organic farming, Organic apples are commonly produced in the United States. Due to infestations by key insects and diseases, organic production is difficult in Europe. 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. A light coating of kaolin, which forms a physical barrier to some pests, also may help prevent apple sun scalding.


Phytochemicals

Apple skins and seeds contain various phytochemicals, particularly polyphenols which are under preliminary research for their potential health effects.


Non-browning apples

The enzyme polyphenol oxidase causes Food browning, browning in sliced or bruised apples, by catalysis, catalyzing the redox, oxidation of phenolic compounds to o-quinones, a browning factor. Browning reduces apple taste, color, and food value. Arctic Apples, Arctic apples, a non-browning group of apples introduced to the United States market in 2019, have been Genetically modified food, genetically modified to silence the gene expression, expression of polyphenol oxidase, thereby delaying a browning effect and improving apple eating quality. The US Food and Drug Administration in 2015, and Canadian Food Inspection Agency in 2017, determined that Arctic apples are as safe and nutritious as conventional apples.


Other products

Apple seed oil is obtained by Expeller pressing, pressing apple seeds for manufacturing cosmetics.


Research

Preliminary research is investigating whether apple consumption may affect the risk of some types of cancer.


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 birch pollen. 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, but in rare cases can also include life-threatening anaphylaxis. 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. In other areas, such as the Mediterranean, some individuals have adverse reactions to apples because of their similarity to peaches. 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. Breeding efforts have yet to produce a hypoallergenic fruit suitable for either of the two forms of apple allergy.


Toxicity of seeds

Apple seeds contain small amounts of amygdalin, a sugar and cyanide compound known as a Cyanogenic glycoside#Cyanogenic glycosides, 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. The United States National Library of Medicine's Hazardous Substances Data Bank records no cases of amygdalin poisoning from consuming apple seeds.


See also

* Apple chip * Applecrab, apple–crabapple hybrids for eating * Cooking apple * Johnny Appleseed * List of apple cultivars * List of apple dishes * Rootstock * Welsh apples


References


Further reading

Books * * *


External links

* {{Authority control Apples, * Malus, * Fruits originating in Asia Plants described in 1768