{{taxobox |name = Apple |image = Red Apple.jpg |image_caption = A typical apple |regnum = Plantae |unranked_divisio = Angiosperms |unranked_classis = Eudicots |unranked_ordo = Rosids |ordo = Rosales |familia = Rosaceae |subfamilia = Maloideae or Spiraeoideae |tribus = Maleae |genus = ''Malus'' |species = ''M. domestica'' |binomial = ''Malus domestica'' |binomial_authority = Borkh. |}}

The apple is the pomaceous fruit of the apple tree, species ''Malus domestica'' in the rose family (Rosaceae). It is one of the most widely cultivated tree fruits, and the most widely known of the many members of genus ''Malus'' that are used by humans. Apples grow on small, deciduous trees. The tree originated in Western 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 been present in the mythology and religions of many cultures, including Norse, Greek and Christian traditions. In 2010, the fruit's genome was decoded, leading to new understandings of disease control and selective breeding in apple production.

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 uses, including in cooking, fresh eating and cider production. Domestic apples are generally propagated by grafting, although wild apples grow readily from seed. Trees are prone to a number of fungal, bacterial and pest problems, which can be controlled by a number of organic and non-organic means.

At least 55 million tonnes of apples were grown worldwide in 2005, with a value of about $10 billion. China produced about 35% of this total. The United States is the second-leading producer, with more than 7.5% of world production. Iran is third, followed by Turkey, Russia, Italy and India. Apples are often eaten raw, but can also be found in many foods (especially desserts) and drinks. Many beneficial health effects have been found from eating apples; however, the seeds are slightly poisonous and two forms of allergies are seen to various proteins found in the fruit.

Botanical information

The apple forms a tree that is small and deciduous, reaching tall, with a broad, often densely twiggy crown. The leaves are alternately arranged simple ovals 5 to 12 cm long and broad on a [[Petiole (botany)|petiole ]] with an acute tip, serrated margin and a slightly downy underside. Blossoms are produced in spring simultaneously with the budding of the leaves. The flowers are white with a pink tinge that gradually fades, five petaled, and in diameter. The fruit matures in autumn, and is typically in diameter. The center of the fruit contains five carpels arranged in a five-point star, each carpel containing one to three seeds, called pips.

Wild ancestors

The wild ancestors of ''Malus domestica'' are ''Malus sieversii'', found growing wild in the mountains of Central Asia in southern Kazakhstan, Kyrgyzstan, Tajikistan, and Xinjiang, China,, using the Golden delicious variety. It had about 57,000 genes, the highest number of any plant genome studied to date and more genes than the human genome (about 30,000). This new understanding of the apple genome will help scientists in identifying genes and gene variants that contribute to resistance to disease and drought, and other desirable characteristics. Understanding the genes behind these characteristics will allow scientists to perform more knowledgeable selective breeding. Decoding the genome 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.

History

The center of diversity of the genus ''Malus'' is in eastern Turkey. The apple tree was perhaps the earliest tree to be cultivated, and its fruits have been improved through selection over thousands of years. Alexander the Great is credited with finding dwarfed apples in Kazakhstan in Asia in 328 BCE; those he brought back to Macedonia might have been the progenitors of dwarfing root stocks. Winter apples, picked in late autumn and stored just above freezing, have been an important food in Asia and Europe for millennia, as well as in Argentina and in the United States since the arrival of Europeans. Apples were brought to North America with colonists in the 17th century, and the first apple orchard on the North American continent was said to be near Boston in 1625. In the 20th century, irrigation projects in Washington state began and allowed the development of the multibillion dollar fruit industry, of which the apple is the leading species.

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. In the 21st century, long-term storage again came into popularity, as "controlled atmosphere" facilities were used to keep apples fresh year-round. Controlled atmosphere facilities use high humidity and low oxygen and carbon dioxide levels to maintain fruit freshness.

Cultural aspects

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. 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, and 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 mound. 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.

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 the skald 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 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, but 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 Tree of Life growing at its center.

The Greek goddess of discord, Eris, became disgruntled after she was excluded from the wedding of Peleus and Thetis. In retaliation, she tossed a golden apple inscribed 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.

The apple was thus considered, in ancient Greece, to be sacred to Aphrodite, and 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 (a.k.a. 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.

The forbidden fruit in the Garden of Eden

Though the forbidden fruit 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. This may have been the result of Renaissance painters adding elements of Greek mythology into biblical scenes (alternative interpretations also based on Greek mythology occasionally replace the apple with a pomegranate). In this case the unnamed fruit of Eden became an apple under the influence of 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. In Latin, the words for "apple" and for "evil" are similar (''mālum'' "an apple", ''mălum'' "an evil, a misfortune"). This may also have influenced the apple becoming interpreted as the biblical "forbidden fruit". The larynx in the human throat has been called Adam's apple because of a notion that it was caused by the forbidden fruit sticking in the throat of Adam. The apple as symbol of sexual seduction has been used to imply sexuality between men, possibly in an ironic vein.

Apple cultivars

There are more than 7,500 known cultivars of apples. Cultivars vary in their yield and the ultimate size of the tree, even when grown on the same rootstock. Different cultivars are available for temperate and subtropical climates. One large collection of over 2,100 apple cultivars is housed at the National Fruit Collection in England. 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 flavour that dessert apples cannot.

Commercially popular apple cultivars are soft but crisp. Other desired qualities in modern commercial apple breeding are a colourful skin, absence of russeting, ease of shipping, lengthy storage ability, high yields, disease resistance, typical 'Red Delicious' apple shape, and popular flavour. Modern apples are generally sweeter than older cultivars, as popular tastes in apples have varied over time. Most North Americans and Europeans favour sweet, subacid apples, but tart apples have a strong minority following. Extremely sweet apples with barely any acid flavour are popular in Asia and especially India.

Old cultivars are often oddly shaped, russeted, and have a variety of textures and colours. Some find them to have a better flavour than modern cultivars,

Apple production

Apple breeding

In the wild, apples grow quite readily from seeds. However, like most perennial fruits, apples are ordinarily propagated asexually by grafting. This is because seedling apples are an example of "extreme heterozygotes", in that rather than inheriting DNA from their parents to create a new apple with those characteristics, they are instead different from their parents, sometimes radically. Triploids have an additional reproductive barrier in that the 3 sets of chromosomes cannot be divided evenly during meiosis, yielding unequal segregation of the chromosomes (aneuploids). Even in the very unusual case when a triploid plant can produce a seed (apples are an example), it happens infrequently, and seedlings rarely survive. 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.

The Excelsior Experiment Station of the University of Minnesota has, since the 1930s, introduced a steady progression of important hardy apples that are widely grown, both commercially and by backyard orchardists, throughout Minnesota and Wisconsin. Its most important introductions have included 'Haralson' (which is the most widely cultivated apple in Minnesota), 'Wealthy', 'Honeygold', and 'Honeycrisp'.

Apples have been acclimatized in Ecuador at very high altitudes, where they provide crops twice per year because of constant temperate conditions year-round.

Apple rootstocks

Rootstocks used to control tree size have been used in apple cultivation for over 2,000 years. Dwarfing rootstocks were probably discovered by chance in Asia. Alexander the Great sent samples of dwarf apple trees back to his teacher, Aristotle, in Greece. They were maintained at the Lyceum, a center of learning in Greece.

Most modern apple rootstocks were bred in the 20th century. Much research into the existing rootstocks was begun at the East Malling Research Station in Kent, England. Following that research, Malling worked with the John Innes Institute and Long Ashton to produce a series of different rootstocks with disease resistance and a range of different sizes, which have been used all over the world.

Pollination

Apples are self-incompatible; they must cross-pollinate to develop fruit. During the flowering each season, apple growers usually provide pollinators to carry the pollen. Honey bees are most commonly used. Orchard mason bees are also used as supplemental pollinators in commercial orchards. Bumblebee queens are sometimes present in orchards, but not usually in enough quantity to be significant pollinators.

There are four to seven pollination groups in apples, depending on climate:

Group A – Early flowering, 01 to 3 May in England (Gravenstein, Red Astrachan)

Group B – 04 to 7 May (Idared, McIntosh)

Group C – Mid-season flowering, 08 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).

Varieties are sometimes classed as to the day of peak bloom in the average 30 day blossom period, with pollinizers selected from varieties 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, will grow very large, which allows them to bear much more fruit, but makes harvesting very difficult. 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. Dwarf trees will bear about of fruit per year.

Storage

Commercially, apples can be stored for some months in controlled-atmosphere chambers to delay ethylene-induced onset of ripening. The 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 moving too quickly. Ripening continues when the fruit is removed. For home storage, most varieties of apple can be held for approximately two weeks when kept at the coolest part of the refrigerator (i.e. below 5 °C). Some types, including the Granny Smith and Fuji, can be stored up to a year without significant degradation.

Pests and diseases

The trees are susceptible to a number of fungal and bacterial diseases and insect pests. Many commercial orchards pursue an aggressive program of chemical sprays to maintain high fruit quality, tree health, and high yields. A trend in orchard management is the use of organic methods. These use a less aggressive and direct methods of conventional farming. Instead of spraying potent chemicals, often shown to be potentially dangerous and maleficent to the tree in the long run, organic methods include encouraging or discouraging certain cycles and pests. To control a specific pest, organic growers might encourage the prosperity of its natural predator instead of outright killing it, and with it the natural biochemistry around the tree. Organic apples generally have the same or greater taste than conventionally grown apples, with reduced cosmetic appearances.

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 are fireblight, a bacterial disease; and ''Gymnosporangium'' rust, and black spot, two fungal diseases. Codling moths and apple maggots are two other pests which affect apple trees. 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.

Commerce

At least 55 million tonnes of apples were grown worldwide in 2005, with a value of about $10 billion. About two-fifths of this total was produced in China. More than 7.5% world production occurs in the United States.

In the United States, more than 60% of all the apples sold commercially are grown in Washington state. Imported apples from New Zealand and other more temperate areas are competing with US production and increasing each year.

Most of Australia's apple production is for domestic consumption. Imports from New Zealand have been disallowed under quarantine regulations for fireblight since 1921.

The largest exporters of apples in 2006 were China, Chile, Italy, France and the U.S., while the biggest importers in the same year were Russia, Germany, the UK and the Netherlands.

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+ Top ten apple producers – 11 June 2008	Country		tonnes)	! Footnote
	27 507 000	F
	4 237 730 || align=right |
	2 660 000 || align=right>F
<		2 266 437 || align=right |
	2 211 000 || align=right>F
<		2 072 500 || align=right |
	2 001 400 || align=right |
	1 800 000 || align=right>F
<		1 390 000 || align=right>F
<		1 300 000 || align=right>F
	Source: FAO

Human consumption

Apples are often eaten raw; except for the seeds, which are slightly poisonous (see below), the whole fruit including the skin is suitable for human consumption. Varieties bred for this purpose are termed dessert or table apples.

Apples can be canned or juiced. They are milled to produce apple cider (non-alcoholic, sweet cider) and filtered for apple juice. The juice can be fermented to make cider (alcoholic, hard cider), ciderkin, and vinegar. Through distillation, various alcoholic beverages can be produced, such as applejack, Calvados, and apfelwein. Pectin and apple seed oil may also be produced.

Apples are an important ingredient in many desserts, such as apple pie, apple crumble, apple crisp and apple cake. They are often eaten baked or stewed, and they can also be dried and eaten or reconstituted (soaked in water, alcohol or some other liquid) for later use. Puréed apples are generally known as apple sauce. Apples are also made into apple butter and apple jelly. They are also used (cooked) in meat dishes.

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 US are candy apples (coated in a hard shell of crystallised 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. Farms with apple orchards may open them to the public, so consumers may themselves pick the apples they will buy.

Sliced apples turn brown with exposure to air due to the conversion of natural phenolic substances into melanin upon exposure to oxygen. Different cultivars vary in their propensity to brown after slicing. Sliced fruit can be treated with acidulated water to prevent this effect.

Organic apples are commonly produced in the United States. Organic production is difficult in Europe, though a few orchards have done so with commercial success, using disease-resistant cultivars and the very best cultural controls. The latest tool in the organic repertoire is a spray of a light coating of kaolin clay, which forms a physical barrier to some pests, and also helps prevent apple sun scald.

Apple 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. The allergy is caused by a protein in apples that is similar to birch pollen, and people affected by this protein can also become allergic to other fruits, nuts and vegetables. Reactions, which are called 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, people have adverse reactions to apples because of their similarity to peaches, including a close relationship between the allergens of the two fruits. 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 either 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 for either of the two types of apple allergy.

Nutrition

[[File:Health benefits of apples.png|thumb|left|Health benefits of apple consumption. ]]

The proverb "''An apple a day keeps the doctor away.''", addressing the health effects of the fruit, dates from 19th century Wales. Research suggests that apples may reduce the risk of colon cancer, prostate cancer and lung cancer. Compared to many other fruits and vegetables, apples contain relatively low amounts of vitamin C, but are a rich source of other antioxidant compounds. The fiber content, while less than in most other fruits, helps regulate bowel movements and may thus reduce the risk of colon cancer. They may also help with heart disease, weight loss, and controlling cholesterol. The fiber contained in apples reduces cholesterol by preventing reabsorption, and (like most fruits and vegetables) they are bulky for their caloric content. However, apple seeds are mildly poisonous, containing a small amount of amygdalin, a cyanogenic glycoside. It usually is not enough to be dangerous to humans, but can deter birds.

There is evidence from laboratory experiments that apples possess phenolic compounds which may be cancer-protective and demonstrate antioxidant activity. The predominant phenolic phytochemicals in apples are quercetin, epicatechin, and procyanidin B2.

Apple juice concentrate has been found to increase the production of the neurotransmitter acetylcholine in mice, providing a potential mechanism for the "prevention of the decline in cognitive performance that accompanies dietary and genetic deficiencies and aging." Other studies have shown an "alleviation of oxidative damage and cognitive decline" in mice after the administration of apple juice. Researchers at the Chinese University of Hong Kong discovered that fruit flies who were fed an apple extract lived 10% longer than other flies who were fed a normal diet.

References

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Further reading

;Books

Browning, F. (1999). ''Apples: The Story of the Fruit of Temptation''. North Point Press. ISBN 9780865475793

Mabberley, D.J. / Juniper, B.E. (2009). ''The Story of the Apple''. Timber Press. ISBN 9781604691726

;Review articles on possible health benefits

Gerhauser, C. “Cancer chemopreventive potential of apples, apple juice, and apple components”, Planta Medica (2008),74(13):1608–1624.

Hyson, D.A. “A Comprehensive Review of Apples and Apple Components and Their Relationship to Human Health”, Advances in Nutrition. An International Review Journal (2011),2(5):408–420.

External links

Apple Facts from the UK's Institute of Food Research

National Fruit Collection (UK)

Brogdale Farm (home of the UK's National Fruit Collection)

Grand Valley State University digital collections- diary of Ohio fruit farmer Theodore Peticolas, 1863

Apples Category:Maleae Category:Sequenced genomes

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