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An apple is a round, edible fruit produced by an apple tree ('' Malus'' spp.). Fruit trees of the orchard or domestic apple (''Malus domestica''), the most widely grown in the genus, are cultivated worldwide. The tree originated in
Central Asia Central Asia is a region of Asia consisting of Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan. The countries as a group are also colloquially referred to as the "-stans" as all have names ending with the Persian language, Pers ...
, where its wild ancestor, '' Malus sieversii'', is still found. Apples have been grown for thousands of years in
Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
before they were introduced to
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
by European colonists. Apples have cultural significance in many mythologies (including Norse and Greek) and
religion Religion is a range of social system, social-cultural systems, including designated religious behaviour, behaviors and practices, morals, beliefs, worldviews, religious text, texts, sanctified places, prophecies, ethics in religion, ethics, or ...
s (such as Christianity in Europe). Apples grown from
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s tend to be very different from those of their parents, and the resultant fruit frequently lacks desired characteristics. For commercial purposes, including botanical evaluation, apple cultivars are propagated by clonal grafting onto rootstocks. Apple trees grown without rootstocks tend to be larger and much slower to fruit after planting. Rootstocks are used to control the speed of growth and the size of the resulting tree, allowing for easier harvesting. There are more than 7,500 cultivars of apples. Different cultivars are bred for various tastes and uses, including
cooking Cooking, also known as cookery or professionally as the culinary arts, is the art, science and craft of using heat to make food more palatable, digestible, nutritious, or Food safety, safe. Cooking techniques and ingredients vary widely, from ...
, eating raw, and
cider Cider ( ) is an alcoholic beverage made from the Fermented drink, fermented Apple juice, juice of apples. Cider is widely available in the United Kingdom (particularly in the West Country) and Ireland. The United Kingdom has the world's highest ...
or apple juice production. Trees and fruit are prone to fungal, bacterial, and pest problems, which can be controlled by a number of organic and non-organic means. In 2010, the fruit's genome was sequenced as part of research on disease control and selective breeding in apple production.


Etymology

The word ''apple'', whose
Old English Old English ( or , or ), 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 developed from the languages brought to Great Britain by Anglo-S ...
ancestor is , is descended from the
Proto-Germanic Proto-Germanic (abbreviated PGmc; also called Common Germanic) is the linguistic reconstruction, reconstructed proto-language of the Germanic languages, Germanic branch of the Indo-European languages. Proto-Germanic eventually developed from ...
noun , descended in turn from
Proto-Indo-European Proto-Indo-European (PIE) is the reconstructed common ancestor of the Indo-European language family. No direct record of Proto-Indo-European exists; its proposed features have been derived by linguistic reconstruction from documented Indo-Euro ...
. As late as the 17th century, the word also functioned as a generic term for all fruit, including nuts. This can be compared to the 14th-century
Middle English 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 pe ...
expression , meaning a banana.


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 Leaf, 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. Young twigs are covered in fine downy hairs; they become hairless when older. The buds are egg-shaped and dark red or purple in color; they range in size from 3 to 5 millimeters, but are usually less than 4 mm. The bud scales have very hairy edges. When emerging from the buds, the leaves are , meaning that their edges overlap each other. Leaves can be simple ovals ( elliptic), medium or wide in width, somewhat egg-shaped with the wider portion toward their base ( ovate), or even with sides that are more parallel to each other instead of curved ( oblong) with a narrow pointed end. 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 are produced in spring 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 petals 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 In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...
consisting of a cyme with 3–7 flowers. The central flower of the inflorescence is called the "king bloom"; it opens first and can develop a larger fruit. Open apple blossoms are damaged by even brief exposures to temperatures or less, although the overwintering wood and buds are hardy down to .


Fruit

The fruit is a pome that matures in late summer or autumn. 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.


Chemistry

Important volatile compounds in apples that contribute to their scent and flavour include acetaldehyde, ethyl acetate, 1-butanal,
ethanol Ethanol (also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound with the chemical formula . It is an Alcohol (chemistry), alcohol, with its formula also written as , or EtOH, where Et is the ps ...
, 2-methylbutanal, 3-methylbutanal, ethyl propionate, ethyl 2-methylpropionate, ethyl butyrate, ethyl 2-methyl butyrate, hexanal, 1-butanol, 3-methylbutyl acetate, 2-methylbutyl acetate, 1-propyl butyrate, ethyl pentanoate, amyl acetate, 2-methyl-1-butanol, trans-2-hexenal, ethyl hexanoate, hexanol.


Taxonomy

The apple as a species has more than 100 alternative scientific names, or synonyms. 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 wh ...
: 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, under an alternate taxonomy. When first classified by
Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
in 1753, the pears, apples, and quinces were combined into one genus that he named '' Pyrus'' and he named the apple as ''Pyrus malus''. This was widely accepted, however the botanist Philip Miller published an alternate classification in The Gardeners Dictionary with the apple species separated from ''Pyrus'' in 1754. He did not clearly indicate that by ''Malus pumila'' he meant the domesticated apple. Nonetheless, it was used as such by many botanists. When Moritz Balthasar Borkhausen published his scientific description of the apple in 1803 it may have been a new combination of ''P. malus'' var. ''domestica'', but this was not directly referenced by Borkhausen. The earliest use of var. ''domestica'' for the apple was by Georg Adolf Suckow in 1786.


Genome

Apples are diploid, with two sets of chromosomes per cell (though triploid cultivars, with three sets, are not uncommon), have 17 chromosomes and an estimated genome size of approximately 650 Mb. Several whole genome sequences have been completed and made available. The first one in 2010 was based on the diploid cultivar ' Golden Delicious'. 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. Mos ...
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 is a region of Asia consisting of Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan. The countries as a group are also colloquially referred to as the "-stans" as all have names ending with the Persian language, Pers ...
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'', found growing wild in the mountains of Central Asia in southern
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a landlocked country primarily in Central Asia, with a European Kazakhstan, small portion in Eastern Europe. It borders Russia to the Kazakhstan–Russia border, north and west, China to th ...
,
Kyrgyzstan Kyrgyzstan, officially the Kyrgyz Republic, is a landlocked country in Central Asia lying in the Tian Shan and Pamir Mountains, Pamir mountain ranges. Bishkek is the Capital city, capital and List of cities in Kyrgyzstan, largest city. Kyrgyz ...
,
Tajikistan Tajikistan, officially the Republic of Tajikistan, is a landlocked country in Central Asia. Dushanbe is the capital city, capital and most populous city. Tajikistan borders Afghanistan to the Afghanistan–Tajikistan border, south, Uzbekistan to ...
, and northwestern China. Cultivation of the species, most likely beginning on the forested flanks of the Tian Shan mountains, progressed over a long period of time and permitted secondary introgression of genes from other species into the open-pollinated seeds. Significant exchange with '' Malus sylvestris'', the crabapple, resulted in populations of apples being more related to crabapples than to the more morphologically similar progenitor ''Malus sieversii''. In strains without recent admixture the contribution of the latter predominates. The apple is thought to have been domesticated 4,000–10,000 years ago in the Tian Shan mountains, and then to have travelled along the Silk Road to Europe, with hybridization and introgression of wild crabapples from Siberia (''M. baccata''), the Caucasus (''M. orientalis''), and Europe (''M. sylvestris''). Only the ''M. sieversii'' trees growing on the western side of the Tian Shan mountains contributed genetically to the domesticated apple, not the isolated population on the eastern side. 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 between 6570 and 5684 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 (term originally coined in English language) is a geopolitical region encompassing the Arabian Peninsula, the Levant, Turkey, Egypt, Iran, and Iraq. The term came into widespread usage by the United Kingdom and western Eur ...
. There is direct evidence, apple cores, dated to the 10th century BCE from a Judean site between the Sinai and Negev. 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 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 is the capital and most populous city in the Commonwealth (U.S. state), Commonwealth of Massachusetts in the United States. The city serves as the cultural and Financial centre, financial center of New England, a region of the Northeas ...
by Reverend William Blaxton in 1640. The only apples native to North America are crab apples. Apple cultivars brought as seed from Europe were spread along Native American trade routes, as well as being cultivated on colonial farms. An 1845 United States apples nursery catalogue sold 350 of the "best" cultivars, showing the proliferation of new North American cultivars by the early 19th century. In the 20th century, irrigation projects in Eastern Washington 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 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 and Asia Minor.
Alexander the Great Alexander III of Macedon (; 20/21 July 356 BC – 10/11 June 323 BC), most commonly known as Alexander the Great, was a king of the Ancient Greece, ancient Greek kingdom of Macedonia (ancient kingdom), Macedon. He succeeded his father Philip ...
sent samples of dwarf apple trees to
Aristotle Aristotle (; 384–322 BC) was an Ancient Greek philosophy, Ancient Greek philosopher and polymath. His writings cover a broad range of subjects spanning the natural sciences, philosophy, linguistics, economics, politics, psychology, a ...
'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 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. 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

File:Apple tree blossom.JPG, An apple blossom from an old Ayrshire cultivar File:Orchmason.jpg, An orchard mason bee on an apple bloom Apples are self-incompatible; they must cross-pollinate 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 carpel, 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 ...
s to carry 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 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', ' McIntosh') * Group C – Mid-season flowering, 8 to 11 May (' Granny Smith', ' Cox's Orange Pippin') * Group D – Mid/late season flowering, 12 to 15 May (' Golden Delicious', 'Calville blanc d'hiver') * Group E – Late flowering, 16 to 18 May (' Braeburn', 'Reinette d'Orléans') * Group F – 19 to 23 May ('Suntan') * Group H – 24 to 28 May ('Court-Pendu Gris' – also called Court-Pendu plat) One cultivar can be pollinated by a compatible cultivar from the same group or close (A with A, or A with B, but not A with C or D).


Maturation and harvest

File:Relander and apples.jpg, Lauri Kristian Relander, the former President of Finland, with his family picking apples in the 1930s File:SweeTango apples harvested 2010.jpg, Apples being harvested in Wenatchee, Washington, United States (2010) 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 ' Sweet Bough' and 'Duchess'; fall producers include 'Blenheim'; winter producers include '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 Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
, which also must be controlled. For home storage, most culitvars of apple can be stored for three weeks in a pantry and four to six weeks from the date of purchase in a refrigerator that maintains . Some varieties of apples (e.g. ' Granny Smith' and ' 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 and bacterial diseases, and to damage by insect pests. Many commercial orchards pursue a program of chemical sprays to maintain high fruit quality, tree health, and high yields. These prohibit the use of synthetic pesticides, though some older pesticides are allowed. Organic methods include, for instance, introducing its natural predator to reduce the population of a particular pest. A wide range of pests and diseases can affect the plant. Three of the more common diseases or pests are mildew, aphids, and apple scab. * Mildew is characterized by light grey powdery patches appearing on the leaves, shoots and flowers, normally in spring. The flowers turn a creamy yellow color and do not develop correctly. This can be treated similarly to ''Botrytis''—eliminating the conditions that caused the disease and burning the infected plants are among recommended actions. * Aphids are small insects with sucking mouthparts. Five species of aphids commonly attack apples: apple grain aphid, rosy apple aphid, apple aphid, spirea aphid, and the woolly apple aphid. The aphid species can be identified by color, time of year, and by differences in the cornicles (small paired projections from their rear). 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, and 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 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 cultivars (cultivated varieties) 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 In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
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 drink, fermented Apple juice, juice of apples. Cider is widely available in the United Kingdom (particularly in the West Country) and Ireland. The United Kingdom has the world's highest ...
. Cider apples are typically too tart and astringent to eat fresh, but they give the beverage a rich flavor that dessert apples cannot. In Europe, apple breeding programs are conducted at places such as Julius Kühn-Institut, the German federal research center for cultivated plants. In the United States there are many apple breeding programs associated with universities. For instance, in the East, Cornell University has had a program operating since 1880 in Geneva, New York, while in the West, Washington State University started a program to support their home state's apple industry in 1994. Released by the
University of Minnesota The University of Minnesota Twin Cities (historically known as University of Minnesota) is a public university, public Land-grant university, land-grant research university in the Minneapolis–Saint Paul, Twin Cities of Minneapolis and Saint ...
in 1991, the ' Honeycrisp' has become famous for its crispness and juiciness, thereby commanding high market prices. Unusually for a popular cultivar, the 'Honeycrisp' is not directly related to another popular apple cultivar but instead to two unsuccessful cultivars. However, it is also difficult to grow and to store, prompting the industry to seek hybrids that not only appeal to consumers are also less costly for farmers to cultivate and last longer in storage. By the 2020s, about half of the new apple varieties entering the market in the United States and Canada are 'Honeycrisp' progeny. Such hybrids include the ' SweeTango' (a cross between the 'Honeycrisp' and the ' Zestar') introduced by the University of Minnesota in 2008 and the ' Cosmic Crisp' (the 'Honeycrisp' and the ' Enterprise') released by Washington State University in 2017. Commercially popular apple cultivars are soft but crisp. Other desirable qualities in modern commercial apple breeding are a colorful skin, absence of russeting, ease of shipping, lengthy storage ability, high yields, disease resistance, common apple shape, and developed flavor. Modern apples are generally sweeter than older cultivars, as popular tastes in apples have varied over time. Most North Americans and Europeans favor crunchy, sweet, and subacid apples. Nevertheless, tart apples maintain a strong minority following. In the United States today, the most popular apple varieties are the '
Ambrosia In the ancient Greek mythology, Greek myths, ambrosia (, ) is the food or drink of the Greek gods, and is often depicted as conferring longevity or immortality upon whoever consumed it. It was brought to the gods in Mount Olympus, Olympus by do ...
', 'Honeycrisp', and '
Jazz Jazz is a music genre that originated in the African-American communities of New Orleans, Louisiana, in the late 19th and early 20th centuries. Its roots are in blues, ragtime, European harmony, African rhythmic rituals, spirituals, h ...
', according Nielsen data, while in Canada, the 'Honeyscrisp', 'Ambrosia', and 'Gala' take the top spots. Together, these relatively newer varieties have overtaken once dominant cultivars like the ' McIntosh' and the ' Red Delicious' in the North American market. Extremely sweet apples with barely any acid flavor are popular in Asia, especially the
Indian subcontinent The Indian subcontinent is a physiographic region of Asia below the Himalayas which projects into the Indian Ocean between the Bay of Bengal to the east and the Arabian Sea to the west. It is now divided between Bangladesh, India, and Pakista ...
. Old cultivars are often oddly shaped, russeted, and grow in a variety of textures and colors. Some find them to have better flavor than modern cultivars, but they may have other problems that make them commercially unviable—low yield, disease susceptibility, poor tolerance for storage or transport, or just being the "wrong" size. A few old cultivars are still produced on a large scale, but many have been preserved by home gardeners and farmers that sell directly to local markets. Many unusual and locally important cultivars with their own unique taste and appearance exist; apple conservation campaigns have sprung up around the world to preserve such local cultivars from extinction. In the United Kingdom, old cultivars such as ' Cox's Orange Pippin' and ' Egremont Russet' are still commercially important even though by modern standards they are low yielding and susceptible to disease.


Production

World production of apples in 2023 was 97 million tonnes, with China producing 51% of the total (table). Secondary producers were the United States,
Turkey Turkey, officially the Republic of Türkiye, is a country mainly located in Anatolia in West Asia, with a relatively small part called East Thrace in Southeast Europe. It borders the Black Sea to the north; Georgia (country), Georgia, Armen ...
, and
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It extends from the Baltic Sea in the north to the Sudetes and Carpathian Mountains in the south, bordered by Lithuania and Russia to the northeast, Belarus and Ukrai ...
.


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 Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
. 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 Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
(table). A reference serving of a raw apple with skin weighing provides 52 calories and a moderate content of dietary fiber (table). Otherwise, there is low content of micronutrients, with the Daily Values of all falling below 10% (table).


Culinary

Apples varieties can be grouped as cooking apples, eating apples, and cider apples, the last so astringent as to be "almost inedible". Apples are consumed as juice, raw in salads, baked in pies, cooked into 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 Normandy (; or ) is a geographical and cultural region in northwestern Europe, roughly coextensive with the historical Duchy of Normandy. Normandy comprises Normandy (administrative region), mainland Normandy (a part of France) and insular N ...
. 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 Rosh Hashanah (, , ) is the New Year in Judaism. The Hebrew Bible, biblical name for this holiday is Yom Teruah (, , ). It is the first of the High Holy Days (, , 'Days of Awe"), as specified by Leviticus 23:23–25, that occur in the late summe ...
. Apples are an important ingredient in many desserts, such as pies, crumbles, and cakes. When cooked, some apple cultivars easily form a puree known as apple sauce, which can be cooked down to form a preserve, apple butter. They are often baked or stewed, and are cooked in some meat dishes. Apples are milled or pressed to produce apple juice, which may be drunk unfiltered (called apple cider in North America), or filtered. Filtered juice is often concentrated and frozen, then reconstituted later and consumed. Apple juice can be fermented to make
cider Cider ( ) is an alcoholic beverage made from the Fermented drink, fermented Apple juice, juice of apples. Cider is widely available in the United Kingdom (particularly in the West Country) and Ireland. The United Kingdom has the world's highest ...
(called hard cider in North America), ciderkin, and vinegar. Through distillation, various alcoholic beverages can be produced, such as applejack, Calvados, and apple brandy.


Organic production

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
polyphenol Polyphenols () are a large family of naturally occurring phenols. They are abundant in plants and structurally diverse. Polyphenols include phenolic acids, flavonoids, tannic acid, and ellagitannin, some of which have been used historically as ...
s. These are oxidised by the
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
polyphenol oxidase, which causes browning in sliced or bruised apples, by 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 to silence the expression of polyphenol oxidase, thereby delaying a browning effect and improving apple eating quality. The US
Food and Drug Administration The United States Food and Drug Administration (FDA or US FDA) is a List of United States federal agencies, federal agency of the United States Department of Health and Human Services, Department of Health and Human Services. The FDA is respo ...
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 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 Germanic paganism or Germanic religion refers to the traditional, culturally significant religion of the Germanic peoples. With a chronological dating, chronological range of at least one thousand years in an area covering Scandinavia, the Bri ...
, from which
Norse paganism Old Norse religion, also known as Norse paganism, is a branch of Germanic paganism, Germanic religion which developed during the Proto-Norse language, Proto-Norse period, when the North Germanic peoples separated into Germanic peoples, distinc ...
developed. She points out that buckets of apples were found in the Oseberg ship burial site in Norway, and that fruits and nuts have been found in the early graves of the
Germanic peoples The Germanic peoples were tribal groups who lived in Northern Europe in Classical antiquity and the Early Middle Ages. In modern scholarship, they typically include not only the Roman-era ''Germani'' who lived in both ''Germania'' and parts of ...
in England and elsewhere in Europe. The fruits and nuts may have had a symbolic meaning, and 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 Frigg (; Old Norse: ) is a goddess, one of the Æsir, in Germanic mythology. In Norse mythology, the source of most surviving information about her, she is associated with marriage, prophecy, clairvoyance and motherhood, and dwells in the wetl ...
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 A mound is a wikt:heaped, heaped pile of soil, earth, gravel, sand, rock (geology), rocks, or debris. Most commonly, mounds are earthen formations such as hills and mountains, particularly if they appear artificial. A mound may be any rounded ...
. 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" 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 The Roman Empire ruled the Mediterranean and much of Europe, Western Asia and North Africa. The Roman people, Romans conquered most of this during the Roman Republic, Republic, and it was ruled by emperors following Octavian's assumption of ...
and came to Europe from the
Near East The Near East () is a transcontinental region around the Eastern Mediterranean encompassing the historical Fertile Crescent, the Levant, Anatolia, Egypt, Mesopotamia, and coastal areas of the Arabian Peninsula. The term was invented in the 20th ...
, 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, including Greek and Roman mythology where it has an ambiguous symbolism of discord, fertility, or courtship. In
Greek mythology Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of ancient Greek folklore, today absorbed alongside Roman mythology into the broader designation of classical mythology. These stories conc ...
, the Greek hero
Heracles Heracles ( ; ), born Alcaeus (, ''Alkaios'') or Alcides (, ''Alkeidēs''), was a Divinity, divine hero in Greek mythology, the son of ZeusApollodorus1.9.16/ref> and Alcmene, and the foster son of Amphitryon.By his adoptive descent through ...
, 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 The tree of life is a fundamental archetype in many of the world's mythology, mythological, religion, religious, and philosophy, philosophical traditions. It is closely related to the concept of the sacred tree.Giovino, Mariana (2007). ''The ...
growing at its center. The Greek goddess of discord, Eris, became disgruntled after she was excluded from the wedding of Peleus and
Thetis Thetis ( , or ; ) is a figure from Greek mythology with varying mythological roles. She mainly appears as a sea nymph, a goddess of water, and one of the 50 Nereids, daughters of the ancient sea god Nereus. When described as a Nereid in Cl ...
. In retaliation, she tossed a golden apple inscribed Καλλίστη (''Kallistē'', "For the most beautiful one"), into the wedding party. Three goddesses claimed the apple: Hera,
Athena Athena or Athene, often given the epithet Pallas, is an ancient Greek religion, ancient Greek goddess associated with wisdom, warfare, and handicraft who was later syncretism, syncretized with the Roman goddess Minerva. Athena was regarde ...
, and Aphrodite.
Paris Paris () is the Capital city, capital and List of communes in France with over 20,000 inhabitants, largest city of France. With an estimated population of 2,048,472 residents in January 2025 in an area of more than , Paris is the List of ci ...
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, 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 ''Ynys Afallon'', "Island of Apples".


China

In China, apples symbolise peace, since the sounds of the first element ("píng") in the words "apple" (苹果, ''Píngguǒ'') and "peace" (平安, ''Píng'ān'') are homophonous in Mandarin and Cantonese. When these two words are combined, the word ''Píngānguǒ'' (平安果, "Peace apples") is formed. This association developed further as the name for Christmas Eve in Mandarin is ''Píngānyè'' (平安夜, "Peaceful/Quiet Evening"), which made the gifting of apples at this season to friends and associates popular, as a way to wish them peace and safety.


Christian art

Though the forbidden fruit of Eden in the
Book of Genesis The Book of Genesis (from Greek language, Greek ; ; ) is the first book of the Hebrew Bible and the Christian Old Testament. Its Hebrew name is the same as its incipit, first word, (In the beginning (phrase), 'In the beginning'). Genesis purpor ...
is not identified, popular Christian tradition has held that it was an apple that Eve coaxed
Adam Adam is the name given in Genesis 1–5 to the first human. Adam is the first human-being aware of God, and features as such in various belief systems (including Judaism, Christianity, Gnosticism and Islam). According to Christianity, 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 Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
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 The Renaissance ( , ) is a Periodization, period of history and a European cultural movement covering the 15th and 16th centuries. It marked the transition from the Middle Ages to modernity and was characterized by an effort to revive and sur ...
painters may also have been influenced by the story of the golden apples in the Garden of Hesperides. As a result, in the story of Adam and Eve, the apple became a symbol for knowledge, immortality, temptation, the fall of man into sin, and sin itself. The larynx in the human throat has been called the " Adam's apple" because of a notion that it was caused by the forbidden fruit remaining in the throat of Adam. The apple as symbol of sexual seduction has been used to imply human sexuality, possibly in an ironic vein.


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 Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
, 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 * Johnny Appleseed


References


Further reading

* * * * *


External links

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