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An apple is a round, 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 spp.'', among them the domestic or orchard apple; ''Malus domestica''). Apple trees 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 extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''
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 of th ...
''. 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 as ...
'', is still found. Apples have been grown for thousands of years in Eurasia and were introduced to North America by European colonists. Apples have religious and
mythological 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 narrat ...
significance in many cultures, including Norse, Greek, 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. For commercial purposes, including botanical evaluation, 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 Grafting or graftage is a horticultural technique whereby tissues of plants are joined so as to continue their growth together. The upper part of the combined plant is called the scion () while the lower part is called the rootstock. The succ ...
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 Cooking, cookery, or culinary arts is the art, science and craft of using heat to Outline of food preparation, prepare food for consumption. Cooking techniques and ingredients vary widely, from grilling food over an open fire to using electric ...
, eating raw, and
cider Cider ( ) is an alcoholic beverage made from the fermented juice of apples. Cider is widely available in the United Kingdom (particularly in the West Country) and the Republic of Ireland. The UK has the world's highest per capita consumption, ...
or apple juice production. Trees and fruit are prone to
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 Organic may refer to: * Organic, of or relating to an organism, a living entity * Organic, of or relating to an anatomical organ Chemistry * Organic matter, matter that has come from a once-living organism, is capable of decay or is the product ...
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 ge ...
was sequenced as part of research on disease control and selective breeding in apple production.


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 settlement of Britain, Anglo ...
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 branc ...
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-E ...
. 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 Com ...
s. This can be compared to 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 p ...
expression , meaning a
banana A banana is an elongated, edible fruit – botanically a berry – produced by several kinds of large herbaceous flowering plants in the genus ''Musa''. In some countries, bananas used for cooking may be called "plantains", distinguis ...
.


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, aft ...
tree, generally standing tall in cultivation and up to in the wild, though more typically . When cultivated, the size, shape and branch density are determined by rootstock selection and trimming method. Apple trees may naturally have a rounded to erect crown with a dense canopy of leaves. The bark of the trunk is dark gray or gray-brown, but young branches are reddish or dark-brown with a smooth texture. When young twigs are covered in very fine downy hairs and become hairless as they become older. The buds are egg shaped and dark red or purple in color and range in size from 3 to 5 millimeters, but usually less than 4 mm. The
bud scale In botany, a bud is an undeveloped or embryonic shoot and normally occurs in the axil of a leaf or at the tip of a stem. Once formed, a bud may remain for some time in a dormant condition, or it may form a shoot immediately. Buds may be spec ...
s have very hairy edges. When emerging from the buds the leaves are , that is their edges overlap each other. The shape is ranges from simple ovals ( elliptic) medium or wide in width, somewhat egg shaped with the wider portion toward their base (
ovate Ovate may refer to: *Ovate (egg-shaped) leaves, tepals, or other botanical parts *Ovate, a type of prehistoric stone hand axe *Ovates, one of three ranks of membership in the Welsh Gorsedd *Vates In modern English, the nouns vates () and ovat ...
) or even with sides that are more parallel to each other instead of curved (
oblong An oblong is a non-square rectangle. Oblong may also refer to: Places * Oblong, Illinois, a village in the United States * Oblong Township, Crawford County, Illinois, United States * A strip of land on the New York-Connecticut border in the Unit ...
) with a narrow pointed end. The edges have broadly angled teeth, but do not have lobes. The top surface of the leaves are , almost hairless, while the undersides are densely covered in fine hairs. The leaves are attached alternately by short leaf stems long.
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), a season of the year * Spring (device), a mechanical device that stores energy * Spring (hydrology), a natural source of water * Spring (mathematics), a geometric surface in the shape of a ...
simultaneously with the budding of the leaves and are produced on spurs and some long shoots. When the flower buds first begin to open the
petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...
s are rose-pink and fade to white or light pink when fully open with each flower in diameter. The five petaled flowers are group in an
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
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.


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 In botany, a pome is a type of fruit produced by flowering plants in the subtribe Malinae of the family Rosaceae. Well-known pomes include the apple, pear, and quince. Etymology The word ''pome'' entered English in the late 14th century, and re ...
that matures in late
summer Summer is the hottest of the four temperate seasons, occurring after spring and before autumn. At or centred on the summer solstice, the earliest sunrise and latest sunset occurs, daylight hours are longest and dark hours are shortest, wit ...
or
autumn Autumn, also known as fall in American English and Canadian English, is one of the four temperate seasons on Earth. Outside the tropics, autumn marks the transition from summer to winter, in September ( Northern Hemisphere) or March ( Sou ...
. 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. The edible flesh is formed from the receptacle at the base of the flower. 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. The five sepals remain attached and stand out from the surface of the apple. The size of the fruit varies widely between cultivars, but generally has a diameter between . The shape is quite variable and may be nearly round, elongated, conical, or short and wide. 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%. The skin may be wholly or partly russeted, making it rough and brown. The skin is covered in a protective layer of epicuticular wax. The skin may also be marked with scattered dots. The flesh is generally pale yellowish-white, though it can be pink, yellow or green. File:Yellow Transparent (cropped).jpg, 0% overcolor File:Lobo (apple) (cropped).jpg, 100% overcolor


Chemistry

Important volatile compounds in apples that contribute to their scent and flavour include
acetaldehyde Acetaldehyde (IUPAC systematic name ethanal) is an organic chemical compound with the formula CH3 CHO, sometimes abbreviated by chemists as MeCHO (Me = methyl). It is a colorless liquid or gas, boiling near room temperature. It is one of the mos ...
, ethyl acetate, 1-butanal,
ethanol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an Alcohol (chemistry), alcohol with the chemical formula . Its formula can be also written as or (an ethyl ...
, 2-methylbutanal,
3-methylbutanal Isovaleraldehyde organic compound, also known as 3-methylbutanal, with the formula (CH3)2CHCH2CHO. It is an aldehyde, a colorless liquid at STP, and found in low concentrations in many types of food. It can be produced commercially and is used as ...
,
ethyl propionate Ethyl propionate is an organic compound with formula C2H5O2CCH2CH3. It is the ethyl ester of propionic acid. It is a colorless volatile liquid with a pineapple-like odor. Some fruits such as kiwis and strawberries contain ethyl propionate in smal ...
, ethyl 2-methylpropionate,
ethyl butyrate Ethyl butyrate, also known as ethyl butanoate, or butyric ether, is an ester with the chemical formula CH3CH2CH2COOCH2CH3. It is soluble in propylene glycol, paraffin oil, and kerosene. It has a fruity odor, similar to pineapple, and is a key in ...
, ethyl 2-methyl butyrate,
hexanal Hexanal, also called hexanaldehyde or caproaldehyde is an alkyl aldehyde used in the flavor industry to produce fruity flavors. Its scent resembles freshly cut grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monoc ...
,
1-butanol 1-Butanol, also known as butan-1-ol or ''n''-butanol, is a primary alcohol with the chemical formula C4H9OH and a linear structure. Isomers of 1-butanol are isobutanol, butan-2-ol and ''tert''-butanol. The unmodified term butanol usually refers t ...
,
3-methylbutyl acetate Isoamyl acetate, also known as isopentyl acetate, is an organic compound that is the ester formed from isoamyl alcohol and acetic acid, with the molecular formula C7H14O2.It is a colorless liquid that is only slightly soluble in water, but very ...
, 2-methylbutyl acetate, 1-propyl butyrate,
ethyl pentanoate Ethyl pentanoate, also commonly known as ethyl valerate, is an organic compound used in flavors. It is an ester In chemistry, an ester is a compound derived from an oxoacid (organic or inorganic) in which at least one hydroxyl group () is r ...
, amyl acetate,
2-methyl-1-butanol 2-Methyl-1-butanol (IUPAC name, also called active amyl alcohol) is an organic compound with the formula CH3CH2CH(CH3)CH2OH. It is one of several isomers of amyl alcohol. A colorless liquid, it occurs naturally in trace amounts and has attracted ...
, trans-2-hexenal,
ethyl hexanoate Ethyl hexanoate is the ester resulting from the condensation of hexanoic acid and ethanol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an Alcohol (chemis ...
, hexanol.


Taxonomy

The apple as a species has more than 100 alternative scientific names, or
synonyms A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
. 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 A conserved name or ''nomen conservandum'' (plural ''nomina conservanda'', abbreviated as ''nom. cons.'') is a scientific name that has specific nomenclatural protection. That is, the name is retained, even though it violates one or more rules whic ...
: 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. The General Committee of the IAPT decided in June 2017 to approve this change, officially conserving ''M. domestica''. Nevertheless, some works published after 2017 still use ''M. pumila'' as the
correct name In botany, the correct name according to the International Code of Nomenclature for algae, fungi, and plants (ICN) is the one and only botanical name that is to be used for a particular taxon, when that taxon has a particular circumscription, posit ...
, under an alternate taxonomy. When first classified by Linnaeus in 1753, the pears, apples, and quinces were combined into one genus that he named ''
Pyrus Pears are fruits produced and consumed around the world, growing on a tree and harvested in the Northern Hemisphere in late summer into October. The pear tree and shrub are a species of genus ''Pyrus'' , in the family Rosaceae, bearing the po ...
'' 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 ''The Gardeners Dictionary'' was a widely cited reference series, written by Philip Miller (1691–1771), which tended to focus on plants cultivated in England. Eight editions of the series were published in his lifetime. After his death, it was ...
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 Moritz Balthasar Borkhausen (3 December 1760, Giessen – 30 November 1806, Darmstadt) was a German naturalist and forester. He took part in the production of ' by Johann Conrad Susemihl. He received his education in Giessen, and in 1796 starte ...
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. The earliest use of var. ''domestica'' for the apple was by Georg Adolf Suckow in 1786.


Genome

Apples are
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, with two sets of
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s per cell (though triploid cultivars, with three sets, 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 ge ...
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 'Golden Delicious' is a cultivar of apple. It is one of the 15 most popular apple cultivars in the United States. It is not closely related to 'Red Delicious'. History Golden Delicious arose from a chance seedling, possibly a hybrid of 'Gri ...
'. However, this first whole genome sequence contained several errors, in part owing to the high degree of
heterozygosity Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
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 estimates between 42,000 and 44,700 protein-coding genes. 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.


Cultivation


History

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 ...
is generally considered the center of origin for apples due to the genetic variability in specimens there. The wild ancestor of ''Malus domestica'' was ''
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 as ...
'', 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 east. ...
,
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. 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 Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Intr ...
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. The apple is thought to have been domesticated 4,000–10,000 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 The Silk Road () was a network of Eurasian trade routes active from the second century BCE until the mid-15th century. Spanning over 6,400 kilometers (4,000 miles), it played a central role in facilitating economic, cultural, political, and reli ...
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 2,000 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 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 Arabian Peninsula, Arabia (including the Arabian Peninsula and Bahrain), Anatolia, Asia Minor (Asian part of Turkey except Hatay Pro ...
. There was substantial apple production in 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. The Roman writer
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic '' ...
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. 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. For longer storage medieval Europeans strung up cored and peeled apples to dry, either whole or sliced into rings. Of the many Old World plants that the Spanish introduced to Chiloé Archipelago 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 named apple cultivar was introduced 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, Lincolns ...
in 1640. The only apples native to North America are
crab apples ''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 plant, native to the temper ...
. 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 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 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.


Breeding

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. 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 breeders when planted as seeds, propagation usually involves
grafting Grafting or graftage is a horticultural technique whereby tissues of plants are joined so as to continue their growth together. The upper part of the combined plant is called the scion () while the lower part is called the rootstock. The succ ...
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 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 Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkmeni ...
and
Asia Minor Anatolia, tr, Anadolu Yarımadası), and the Anatolian plateau, also known as Asia Minor, is a large peninsula in Western Asia and the westernmost protrusion of the Asian continent. It constitutes the major part of modern-day Turkey. The re ...
.
Alexander the Great Alexander III of Macedon ( grc, wikt:Ἀλέξανδρος, Ἀλέξανδρος, Alexandros; 20/21 July 356 BC – 10/11 June 323 BC), commonly known as Alexander the Great, was a king of the Ancient Greece, ancient Greek kingdom of Maced ...
sent samples of dwarf apple trees to
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of phil ...
'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 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. 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 sport In botany, a sport or bud sport, traditionally called ''lusus'', is a part of a plant that shows morphological differences from the rest of the plant. Sports may differ by foliage shape or color, flowers, fruit, or branch structure. The cause is ...
s (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. 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
cross-pollinate Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...
to develop fruit. During the flowering each season, apple growers often utilize
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...
s to carry pollen. Honey bees are most commonly used. Orchard mason bees are also used as supplemental pollinators in commercial orchards.
Bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gener ...
queens Queens is a borough of New York City, coextensive with Queens County, in the U.S. state of New York. Located on Long Island, it is the largest New York City borough by area. It is bordered by the borough of Brooklyn at the western tip of Long ...
are sometimes present in orchards, but not usually in sufficient number to be significant pollinators. 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') * Group B – 4 to 7 May ('
Idared Idared is a type of apple cultivar from Moscow, Idaho, United States. Variety is characterized by a non-uniform skin color. First developed at the University of Idaho Agricultural Experiment Station in 1942; it is a cross between two apple v ...
', ' McIntosh') * Group C – Mid-season flowering, 8 to 11 May (' Granny Smith', '
Cox's Orange Pippin Cox's Orange Pippin, in Britain often referred to simply as Cox, is an apple cultivar first grown in 1825, at Colnbrook in Buckinghamshire, England, by the retired brewer and horticulturist Richard Cox. Though the parentage of the cultivar i ...
') * Group D – Mid/late season flowering, 12 to 15 May ('
Golden Delicious 'Golden Delicious' is a cultivar of apple. It is one of the 15 most popular apple cultivars in the United States. It is not closely related to 'Red Delicious'. History Golden Delicious arose from a chance seedling, possibly a hybrid of 'Gri ...
', '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).


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. Some 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', '
Sweet Bough 'Sweet Bough' is an early ripening cultivar of domesticated apple An apple is an edible fruit produced by an apple tree (''Malus domestica''). Apple fruit tree, trees are agriculture, cultivated worldwide and are the most widely grown spe ...
', and 'Duchess'; fall producers include 'Fuji', 'Jonagold', 'Golden Delicious', 'Red Delicious', and 'Blenheim'; winter producers include 'Granny Smith', 'King', ' Swayzie', and 'Tolman Sweet'.


Storage

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. 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 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
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 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 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 small insects with Insect mouthparts, 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). 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 three fungal diseases: ''Gymnosporangium'' rust, Black Spot (disease), black spot, and Bitter rot of apple, bitter rot. 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. 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.


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. 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 ( ) is an alcoholic beverage made from the fermented juice of apples. Cider is widely available in the United Kingdom (particularly in the West Country) and the Republic of Ireland. The UK has the world's highest per capita consumption, ...
. Cider apples are typically too tart and astringent to eat fresh, but they give the beverage a rich flavor that dessert apples cannot. In the United States there are many apple breeding programs associated with universities. Cornell University has had a program operating since 1880 in Geneva, New York. Among their recent well known apples is the 'SnapDragon' cultivar released in 2013. In the west Washington State University started a program to support their apple industry in 1994 and released the 'Cosmic Crisp' cultivar in 2017. The third most grown apple cultivar in the United States is the 'Honeycrisp', released by the University of Minnesota program in 1991. Unusually for a popular cultivar, the 'Honeycrisp' is not directly related to another popular apple cultivar but instead to two unsuccessful cultivars. In Europe there are also many breeding programs such as the Julius Kühn-Institut, the German federal research center for cultivated plants. 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 Cox's Orange Pippin, in Britain often referred to simply as Cox, is an apple cultivar first grown in 1825, at Colnbrook in Buckinghamshire, England, by the retired brewer and horticulturist Richard Cox. Though the parentage of the cultivar i ...
' and 'Egremont Russet' are still commercially important even though by modern standards they are low yielding and susceptible to disease.


Production

World production of apples in 2022 was 95 million tonnes, with China producing 50% of the total (table). Secondary producers were the United States and Turkey.


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. The U.S. National Library of Medicine's Hazardous Substances Data Bank records no cases of amygdalin poisoning from consuming apple seeds.


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.


Uses


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%.


Culinary

Apples varieties can be grouped as cooking apples, table apple, eating apples, and cider apples, the last so astringent as to be "almost inedible". Apples are consumed as Apple juice, juice, raw in salads, baked in Apple pie, pies, cooked into Apple sauce, sauces and apple butter, or baked. They are sometimes used as an ingredient in savory foods, such as sausage and stuffing. Several techniques are used to preserve apples and apple products. Traditional methods include drying and making apple butter. Juice and cider are produced commercially; cider is a significant industry in regions such as the West of England and Normandy. A toffee apple (UK) or caramel apple (US) is a confection made by coating an apple in hot toffee or caramel candy respectively and allowing it to cool. Apples and honey are a ritual food pairing eaten during the Jewish New Year of Rosh Hashanah. Apples are an important ingredient in many desserts, such as apple pie, pies, crumbles, and apple cake, 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. Apples are Cider mill, milled or Fruit press#Cider press, 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 Cider ( ) is an alcoholic beverage made from the fermented juice of apples. Cider is widely available in the United Kingdom (particularly in the West Country) and the Republic of Ireland. The UK has the world's highest per capita consumption, ...
(called hard cider in North America), ciderkin, and vinegar. Through distillation, various alcoholic beverages can be produced, such as applejack (beverage), applejack, 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.


Non-browning apples

Apple skins and seeds contain polyphenols. These are oxidised by the enzyme polyphenol oxidase, which causes Food browning, browning in sliced or bruised apples, by catalysis, catalyzing the oxidation of phenolic compounds to o-quinones, a browning factor. Browning reduces apple taste, color, and food value. 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.


In culture


Germanic paganism

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. 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 tumulus, mound. 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. Further, Davidson points out the "strange" phrase "Apples of Hel (location), 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."


Greek mythology

Apples appear in many World religions, religious traditions, often as a mystical or forbidden fruit. One of the problems identifying apples in religion, mythology and Folklore, 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, the Greek hero Heracles, as a part of his Twelve Labours, was required to travel to the Garden of the Hesperides and pick the golden apples off the Hesperides#The Garden of the Hesperides, Tree of Life growing at its center. The Greek goddess of discord, Eris (mythology), Eris, became disgruntled after she was excluded from the wedding of Peleus and Thetis. In retaliation, she tossed a Apple of Discord, golden apple inscribed Kallisti, Καλλίστη (''Kallistē'', "For the most beautiful one"), into the wedding party. Three goddesses claimed the apple: Hera, Athena, and Aphrodite. Paris (mythology), 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 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.


Celtic mythology

In Celtic mythology, the otherworld has many names, including ''Emain Ablach'', "Emain of the Apple-trees". A version of this is Avalon in Arthurian legend, or in Welsh language, Welsh ''Ynys Afallon'', "Island of Apples".


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 wordplay with the 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.


See also

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


References


Further reading

* * *


External links

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