An apple is a sweet, edible fruit produced by an apple tree (Malus pumila).

Apple trees are cultivated worldwide and are the most widely grown species in the genus Malus.
The tree originated in Central Asia, where its wild ancestor, Malus sieversii, 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 and mythological significance in many
cultures, including Norse, Greek and European Christian traditions.
Apple trees are large if grown from seed. Generally, apple cultivars are propagated
by grafting onto rootstocks, which control the size of the resulting tree. There are more than
7,500 known cultivars of apples, resulting in a range of desired characteristics. Different cultivars
are bred for various tastes and use, including cooking, eating raw and cider production. Trees
and fruit are prone to a number of fungal, bacterial and pest problems, which can be controlled
by a number of organic and non-organic means. In 2010, the fruit's genome was sequenced as
part of research on disease control and selective breeding in apple production.
Worldwide production of apples in 2017 was 83.1 million tonnes, with China accounting for half
of the total.[3]

Contents

 1Botanical information
o 1.1Wild ancestors
o 1.2Genome
 2History
 3Society and culture
o 3.1Germanic paganism
o 3.2Greek mythology
o 3.3Christian art
 4Cultivars
 5Cultivation
o 5.1Breeding
o 5.2Pollination
o 5.3Maturation and harvest
o 5.4Storage
o 5.5Pests and diseases
 6Production
 7Nutrition
 8Human consumption
o 8.1Popular uses
o 8.2Organic production
o 8.3Phytochemicals
o 8.4Other products
o 8.5Health effects
o 8.6Allergy
o 8.7Toxicity of seeds
 9Proverbs
 10See also
 11References
 12Further reading
 13External links

Botanical information
Blossoms, fruits, and leaves of the apple tree (Malus pumila)

An apple's side, stem end, and interior

The apple is a deciduous tree, generally standing 6 to 15 ft (1.8 to 4.6 m) tall in cultivation and up
to 30 ft (9.1 m) in the wild. When cultivated, the size, shape and branch density are determined
by rootstock selection and trimming method. The leaves are alternately arranged dark green-
colored simple ovals with serrated margins and slightly downy undersides.[4]

Apple blossom
Blossoms are produced in spring simultaneously with the budding of the leaves and are
produced on spurs and some long shoots. The 3 to 4 cm (1.2 to 1.6 in) flowers are white with a
pink tinge that gradually fades, five petaled, with an inflorescence 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.[4][5]
The fruit matures in late summer or autumn, and cultivars exist in a wide range of sizes.
Commercial growers aim to produce an apple that is 2 3⁄4 to 3 1⁄4 in (7.0 to 8.3 cm) in diameter,
due to market preference. Some consumers, especially those in Japan, prefer a larger apple,
while apples below 2 1⁄4 in (5.7 cm) 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, though
many bi- or tri-colored cultivars may be found.[6] 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.[7] The exocarp
(flesh) is generally pale yellowish-white,[6] though pink or yellow exocarps also occur.

Wild ancestors
Main article: Malus sieversii
The original wild ancestor of Malus pumila was Malus sieversii, found growing wild in
the mountains of Central Asia in southern Kazakhstan, Kyrgyzstan, Tajikistan, and Xinjiang,
China.[4][8] Cultivation of the species, most likely beginning on the forested flanks of the Tian
Shan mountains, progressed over a long period of time and permitted secondary introgression of
genes from other species into the open-pollinated seeds. Significant exchange with Malus
sylvestris, the crabapple, resulted in 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.[9][10][11]

Genome
In 2010, an Italian-led consortium announced they had sequenced the complete genome of the
apple in collaboration with horticultural genomicists at Washington State University,[12] using
'Golden Delicious'.[13] It had about 57,000 genes, the highest number of any plant genome studied
to date[14] and more genes than the human genome (about 30,000).[15] This new understanding of
the apple genome will help scientists identify genes and gene variants that contribute to
resistance to disease and drought, and other desirable characteristics. Understanding the genes
behind these characteristics will help scientists perform more knowledgeable selective breeding.
The genome sequence also provided proof that Malus sieversii was the wild ancestor of the
domestic apple—an issue that had been long-debated in the scientific community.[12]

History

Wild Malus sieversii apple in Kazakhstan

The center of diversity of the genus Malus is in eastern present-day Turkey. The apple tree may
have been the earliest tree that humans cultivated,[16] and growers have improved its fruits
through selection over thousands of years. Alexander the Great is credited with finding dwarfed
apples in Kazakhstan in 328 BCE.[4] Winter apples, picked in late autumn and stored just above
freezing, have been an important food in Asia and Europe for millennia.[16]
Of the many Old World plants that the Spanish introduced to Chiloé Archipelago in the 16th
century, apple trees became particularly well adapted.[17]Apples were introduced to North
America by colonists in the 17th century,[4] and the first apple orchard on the North American
continent was planted in Boston by Reverend William Blaxton in 1625.[18] The only apples native
to North America are crab apples, which were once called "common apples".[19]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.[19] 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.[4]
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.[20][21] 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 used in the United States in the 1960s.[22]
Society and culture
Main article: Apple (symbolism)
Germanic paganism

"Brita as Iduna" (1901) by Carl Larsson

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.
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.[23]
Davidson notes a connection between apples and the Vanir, a tribe of gods associated
with fertility in Norse mythology, citing an instance of eleven "golden apples" being given to woo
the beautiful Gerðr by Skírnir, who was acting as messenger for the major Vanir god Freyr in
stanzas 19 and 20 of Skírnismál. Davidson also notes a further connection between fertility and
apples in Norse mythology in chapter 2 of the Völsunga saga when the major
goddess Frigg sends King Rerir an apple after he prays to Odin for a child, Frigg's messenger (in
the guise of a crow) drops the apple in his lap as he sits atop a mound.[24] Rerir's wife's
consumption of the apple results in a six-year pregnancy and the Caesarean section birth of their
son—the hero Völsung.[25]
Further, Davidson points out the "strange" phrase "Apples of Hel" used in an 11th-century poem
by the skald Thorbiorn Brúnarson. She states this may imply that the apple was thought of by
Brúnarson as the food of the dead. Further, Davidson notes that the potentially Germanic
goddess Nehalennia is sometimes depicted with apples and that parallels exist in early Irish
stories. Davidson asserts that while cultivation of the apple in Northern Europe extends back to
at least the time of the Roman Empire and came to Europe from the Near East, the native
varieties of apple trees growing in Northern Europe are small and bitter. Davidson concludes that
in the figure of Iðunn "we must have a dim reflection of an old symbol: that of the guardian
goddess of the life-giving fruit of the other world."[23]

Greek mythology
Heracles with the apple of Hesperides
Apples appear in many religious traditions, often as a mystical or forbidden fruit. One of the
problems identifying apples in religion, mythology 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.[26] 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 Tree of Life growing at its center.[27][28][29]
The Greek goddess of discord, Eris, became disgruntled after she was excluded from the
wedding of Peleus and Thetis.[30] In retaliation, she tossed a golden
appleinscribed Καλλίστη (Kalliste, sometimes transliterated Kallisti, "For the most beautiful one"),
into the wedding party. Three goddesses claimed the apple: Hera, Athena,
and Aphrodite. Paris of Troy was appointed to select the recipient. After being bribed by both
Hera and Athena, Aphrodite tempted him with the most beautiful woman in the
world, Helen of Sparta. He awarded the apple to Aphrodite, thus indirectly causing the Trojan
War.[31]
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:[32]
I throw the apple at you, and if you are willing to love me, take it and share your girlhood with me;
but if your thoughts are what I pray they are not, even then take it, and consider how short-lived
is beauty.

— Plato, Epigram VII

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),[28] 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.[27]

Christian art
Adam and Eve by Albrecht Dürer(1507), showcasing the apple as a symbol of sin.
Though the forbidden fruit of Eden in the Book of Genesis is not identified, popular Christian
tradition has held that it was an apple that Eve coaxed Adam to share with her.[33] The origin of
the popular identification with a fruit unknown in the Middle East in biblical times is found in
confusion between the Latinwords mālum (an apple) and mălum (an evil), each of which is
normally written malum.[34] 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.[35]
Renaissance painters may also have been influenced by the story of the golden apples in
the Garden of Hesperides. As a result, in the story of Adam and Eve, the apple became a symbol
for knowledge, immortality, temptation, the fall of man into sin, and sin itself. The larynx in the
human throat has been called the "Adam's apple" because of a notion that it was caused by the
forbidden fruit remaining in the throat of Adam.[33] The apple as symbol of sexual seduction has
been used to imply human sexuality,