Apple

An apple is a round, edible fruit produced by an apple tree

(Malus spp., among them the domestic or orchard apple; Apple
Malus domestica). 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. Apples have been grown for thousands of
years in Eurasia and were introduced to North America by 'Cripps Pink' apples
European colonists. Apples have religious and mythological
significance in many cultures, including Norse, Greek, and
European Christian tradition.

Apples grown from seed tend to be very different from those of

their parents, and the resultant fruit frequently lacks desired
characteristics. For commercial purposes, including botanical
evaluation, apple cultivars are propagated by clonal grafting
onto rootstocks. Apple trees grown without rootstocks tend to
Flowers
be larger and much slower to fruit after planting. Rootstocks
are used to control the speed of growth and the size of the Scientific classification
resulting tree, allowing for easier harvesting. Kingdom: Plantae
There are more than 7,500 cultivars of apples. Different Clade: Tracheophytes
cultivars are bred for various tastes and uses, including cooking, Clade: Angiosperms
eating raw, and cider or apple juice production. Trees and fruit
Clade: Eudicots
are prone to fungal, bacterial, and pest problems, which can be
controlled by a number of organic and non-organic means. In Clade: Rosids
2010, the fruit's genome was sequenced as part of research on Order: Rosales
disease control and selective breeding in apple production.
Family: Rosaceae
Genus: Malus
Etymology Species: M. domestica

The word apple, whose Old English ancestor is æppel, is Binomial name
descended from the Proto-Germanic noun *aplaz, descended in Malus domestica
turn from Proto-Indo-European *h₂ébōl.[3] As late as the 17th (Suckow) Borkh.
century, the word also functioned as a generic term for all fruit,
Synonyms[1][2]
including nuts. This can be compared to the 14th-century
Middle English expression appel of paradis, meaning a
M. communis Desf., 1768
banana.[4]
 M. pumila Mil.
Description M. frutescens Medik.

The apple is a deciduous tree, generally standing 2 to 4.5 M. paradisiaca (L.) Medikus
metres (6 to 15 feet) tall in cultivation and up to 15 m (49 ft) in M. sylvestris Mil.
the wild, though more typically 2 to 10 m (6.5 to 33 ft).[5][1 ] Pyrus malus L.
When cultivated, the size, shape and branch density are
determined by rootstock selection and trimming method.[5] Pyrus malus var. paradisiaca
Apple trees may naturally have a rounded to erect crown with L.

a dense canopy of leaves.[6] The bark of the trunk is dark gray Pyrus dioica Moench
or gray-brown, but young branches are reddish or dark-brown
with a smooth texture.[1 ][7 ] Young twigs are covered in fine
downy hairs; they become hairless when older.[7 ]

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 convolute, meaning that their edges overlap each other.[1 ] 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.[7 ][1 ] The edges have broadly-angled teeth, but do not have lobes. The top surface
of the leaves are glabrescent, almost hairless, while the undersides are densely covered in fine hairs.[1 ]
The leaves are attached alternately by short leaf stems 1-to-3.5 cm (1 ⁄2 -to-1 1 ⁄2 in) long.[6][1 ]

Blossoms are produced in spring simultaneously with the budding of the leaves and are produced on
spurs and some long shoots.[5] 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 3-to-4-centimeter (1-to-1 1 ⁄2 -inch) in
diameter.[1 ] The five-petaled flowers are group in an inflorescence consisting of a cyme with 3–7
flowers.[8] The central flower of the inflorescence is called the "king bloom"; it opens first and can
develop a larger fruit.[6] Open apple blossoms are damaged by even brief exposures to temperatures
−2 °C (28 °F) or less, although the overwintering wood and buds are hardy down to −40 °C
(−40 °F).[8]

Apple blossoms Botanical illustration

Fruit
The fruit is a pome that matures in late summer or autumn.[1 ] 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.[9] The edible flesh is
formed from the receptacle at the base of the flower.[1 0]

How apple fruit derives from flower structures

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.[1 1 ] The five
sepals remain attached and stand out from the surface of the apple.[1 ]

The size of the fruit varies widely between cultivars, but generally has a diameter between 2.5 and
12 cm (1 and 5 in).[7 ] The shape is quite variable and may be nearly round, elongated, conical, or short
and wide.[1 2]

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%.[1 3] The skin may be wholly or partly russeted, making it rough and brown. The skin is covered
in a protective layer of epicuticular wax.[1 4] The skin may also be marked with scattered dots.[1 ] The
flesh is generally pale yellowish-white, though it can be pink, yellow or green.[1 3]

Apples can have any amount of overcolor, a darker tint over a pale groundcolor.
 0% overcolor 100% overcolor

Chemistry
Important volatile compounds in apples that contribute to their scent and flavour include
acetaldehyde, ethyl acetate, 1-butanal, ethanol, 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.[1 5][1 6]

Taxonomy
The apple as a species has more than 100 alternative scientific names, or synonyms.[1 7 ] 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:
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.[1 8] The General Committee of the IAPT decided in June 2017 to approve this change,
officially conserving M. domestica.[1 9] Nevertheless, some works published after 2017 still use
M. pumila as the correct name, under an alternate taxonomy.[2]

When first classified by Linnaeus 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.[1 7 ] The
earliest use of var. domestica for the apple was by Georg Adolf Suckow in 1786.[2]

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'.[20] However, this first whole genome sequence
contained several errors,[21 ] in part owing to the high degree of heterozygosity in diploid apples which,
in combination with an ancient genome duplication, complicated the assembly. Recently, double- and
trihaploid individuals have been sequenced, yielding whole genome sequences of higher quality.[22][23]
The first whole genome assembly was estimated to contain around 57,000 genes,[20] though the more
recent genome sequences support estimates between 42,000 and 44,700 protein-coding
genes.[22][23] 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.[24]

Cultivation

History

Map of the origins of the cultivated apple. The wild origin is in Kazakhstan; hybridisations and
repeated domestications followed, modifying many attributes of the fruit.[24]

Central Asia is generally considered the center of origin for apples

due to the genetic variability in specimens there.[25] The wild
ancestor of Malus domestica was Malus sieversii, found growing
wild in the mountains of Central Asia in southern Kazakhstan,
Kyrgyzstan, Tajikistan, and northwestern China.[5][26] 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 Wild Malus sieversii apple in
sylvestris, the crabapple, resulted in populations of apples being Kazakhstan
more related to crabapples than to the more morphologically
similar progenitor Malus sieversii. In strains without recent
admixture the contribution of the latter predominates.[27 ][28][29]
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.[24]

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.[24]

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.[24]

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.[30] 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.[31 ]

There is indirect evidence of apple cultivation in the third millennium BCE in the Middle East.[31 ]
There is direct evidence, apple cores, dated to the 10th century BCE from a Judean site between the
Sinai and Negev. [32] There was substantial apple production in European classical antiquity, and
grafting was certainly known then.[31 ] 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.[31 ]

The Roman writer Pliny the Elder describes a method

"Wild Apples"
of storage for apples from his time in the 1st century. by Henry David Thoreau
He says they should be placed in a room with good air Read by Kev in S for LibriVox
circulation from a north facing window on a bed of 1:01:36
straw, chaff, or mats with windfalls kept
Audio 01:01:35 (full text (https://a
separately.[33] Though methods like this will extend rchive.org/details/excursions1863
the availabity of reasonably fresh apples, without thor/page/266))
refrigeration their lifespan is limited. Even sturdy
winter apple varieties will only keep well until Problems playing this file? See media help.

December in cool climates.[34] For longer storage

medieval Europeans strung up cored and peeled apples to dry, either whole or sliced into rings.[35]

Of the many Old World plants that the Spanish introduced to Chiloé Archipelago in the 16th century,
apple trees became particularly well adapted.[36] Apples were introduced to North America by
colonists in the 17th century,[5] and the first named apple cultivar was introduced in Boston by
Reverend William Blaxton in 1640.[37 ] The only apples native to North America are crab apples.[38]
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.[38] 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.[5]

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.[39][40] 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.[41 ]

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.[42]
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 An apple tree in Germany
(aneuploids). Even in the case when a triploid plant can produce a
seed (apples are an example), it occurs infrequently, and seedlings
rarely survive.[43]

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
3.0 m or 10 ft high at maturity), which bear fruit many years earlier in their life cycle than full size
trees, and are easier to harvest.[44]

Dwarf rootstocks for apple trees can be traced as far back as 300 BCE, to the area of Persia and Asia
Minor. Alexander the Great sent samples of dwarf apple trees to Aristotle's Lyceum. Dwarf rootstocks
became common by the 15th century and later went through several cycles of popularity and decline
throughout the world.[45] 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.[46]

Most new apple cultivars originate as seedlings, which either arise by chance or are bred by
deliberately crossing cultivars with promising characteristics.[47 ] 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.[48]

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.[49]

Pollination

An apple blossom from an old Ayrshire cultivar

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 pollinators 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.[48][50]
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).[51 ]

Maturation and harvest

Lauri Kristian Relander, the former Apples being

President of Finland, with his family harvested in
picking apples in the 1930s 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 40–200 kg (90–440 lb) 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 10–80 kg (20–180 lb) of fruit per year.[48]

Some farms with apple orchards open them to the public so consumers can pick their own apples.[52]

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'.[38]
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 Rome apples on sale at a farmer's
carbon dioxide, which also must be controlled.[53] market in Newark, Delaware, United
States (2011)
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 4 to 0 °C (39 to 32 °F).[54][55] Some varieties of apples (e.g. 'Granny
Smith' and 'Fuji') have more than three times the storage life of others.[56]

Non-organic apples may be sprayed with a substance 1-methylcyclopropene blocking the apples'
ethylene receptors, temporarily preventing them from ripening.[57 ]

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 Codling moth larva tunnelling inside
the more common diseases or pests are mildew, aphids, and apple an 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.[58]
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).[59] 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.[60]
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.[61]
Among the most serious disease problems is a bacterial disease called fireblight, and three fungal
diseases: Gymnosporangium rust, black spot,[62] and bitter rot.[63] 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.[61 ] 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.[64]

Cultivars
There are more than 7,500 known cultivars (cultivated varieties)
of apples.[65] Cultivars vary in their yield and the ultimate size of
the tree, even when grown on the same rootstock.[66] Different
cultivars are available for temperate and subtropical climates. The
UK's National Fruit Collection, which is the responsibility of the
Department of Environment, Food, and Rural Affairs, includes a
collection of over 2,000 cultivars of apple tree in Kent.[67 ] The
University of Reading, which is responsible for developing the UK From left to right: the 'Golden
national collection database, provides access to search the national Delicious', 'SweeTango', 'Granny
collection. The University of Reading's work is part of the Smith', and 'Gala' apples.
European Cooperative Programme for Plant Genetic Resources of
which there are 38 countries participating in the Malus/Pyrus
work group.[68]

The UK's national fruit collection database contains much information on the characteristics and origin
of many apples, including alternative names for what is essentially the same "genetic" apple cultivar.
Most of these cultivars are bred for eating fresh (dessert apples), though some are cultivated
specifically for cooking (cooking apples) or producing cider. Cider apples are typically too tart and
astringent to eat fresh, but they give the beverage a rich flavor that dessert apples cannot.[69]

In Europe, apple breeding programs are conducted at places such as Julius Kühn-Institut, the German
federal research center for cultivated plants.[7 0]

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.[7 1 ] Released by the University of Minnesota in 1991, the 'Honeycrisp' has become
famous for its crispness and juiciness, thereby commanding high market prices.[7 1 ] Unusually for a
popular cultivar, the 'Honeycrisp' is not directly related to another popular apple cultivar but instead
to two unsuccessful cultivars.[7 2] 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.[7 3] By the 2020s, about half of the new apple varieties entering the market
in the United States and Canada are 'Honeycrisp' progeny.[7 4] 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.[7 3]

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.[66] Modern apples are generally
sweeter than older cultivars, as popular tastes in apples have
varied over time.[7 5] Most North Americans and Europeans favor
crunchy, sweet, and subacid apples.[7 6] Nevertheless, tart apples Less common apple cultivars from an
maintain a strong minority following.[7 7 ] In the United States orchard in Italy
today, the most popular apple varieties are the 'Ambrosia',
'Honeycrisp', and 'Jazz', according Nielsen data,[7 3] while in
Canada, the 'Honeyscrisp', 'Ambrosia', and 'Gala' take the top spots.[7 5] Together, these relatively
newer varieties have overtaken once dominant cultivars like the 'McIntosh' and the 'Red Delicious' in
the North American market.[7 5][7 6] Extremely sweet apples with barely any acid flavor are popular in
Asia,[7 7 ] especially the Indian subcontinent.[69]

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.[7 8] 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.[5]

Production
World production of apples in 2023 was 97 million tonnes, with China producing 51% of the total
(table).[7 9] Secondary producers were the United States, Turkey, and Poland.[7 9]

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 Apple production
released. [80] The U.S. National Library of Medicine's Hazardous Substances 2023, millions of
Data Bank records no cases of amygdalin poisoning from consuming apple tonne s
seeds.[81 ]
China 49.6
Allergy United States 5.2
One form of apple allergy, often found in northern Europe, is called birch- Turkey 4.6
apple syndrome and is found in people who are also allergic to birch
Poland 3.9
pollen.[82] Allergic reactions are triggered by a protein in apples that is
similar to birch pollen, and people affected by this protein can also develop India 2.9
allergies to other fruits, nuts, and vegetables. Reactions, which entail oral Iran 2.2
allergy syndrome (OAS), generally involve itching and inflammation of the World 97.3
mouth and throat,[82] but in rare cases can also include life-threatening
Source: FAOSTAT of the United
anaphylaxis.[83] This reaction only occurs when raw fruit is consumed—the
Nations[79]
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.[82]

In other areas, such as the Mediterranean, some individuals have adverse reactions to apples because
of their similarity to peaches.[82] 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.[82]

Breeding efforts have yet to produce a hypoallergenic fruit suitable for either of the two forms of apple
allergy.[82]

Uses

Nutrition
A raw apple is 86% water and 14% carbohydrates, with Apples, with skin (edible parts)
negligible content of fat and protein (table). A reference
serving of a raw apple with skin weighing 100 g (3.5 oz) Nutritional value per 100 g (3.5 oz)
provides 52 calories and a moderate content of dietary Energy 218 kJ (52 kcal)
fiber (table). Otherwise, there is low content of Carbohydrates 13.81 g
micronutrients, with the Daily Values of all falling below Sugars 10.39
10% (table). Dietary fiber 2.4 g
Fat 0.17 g
Protein 0.26 g
Culinary Vitamins and minerals

Vitamins Quantity %DV†

Vitamin A equiv. 3 μg 0%
beta-Carotene 27 μg 0%
lutein zeaxanthin 29 μg
Thiamine (B1) 0.017 mg 1%
Riboflavin (B2) 0.026 mg 2%
Niacin (B3) 0.091 mg 1%
Pantothenic acid (B5) 0.061 mg 1%
Vitamin B6 0.041 mg 2%
Machine for paring, coring, and Folate (B9) 3 μg 1%
slicing apples, from Henry B. Vitamin C 4.6 mg 5%
Scammell's 1897 handbook Vitamin E 0.18 mg 1%
Cyclopedia of Valuable Receipts Vitamin K 2.2 μg 2%
Minerals Quantity %DV†
Apples varieties can be grouped as cooking apples, Calcium 6 mg 0%
eating apples, and cider apples, the last so astringent as Iron 0.12 mg 1%
to be "almost inedible".[86] Apples are consumed as Magnesium 5 mg 1%
juice, raw in salads, baked in pies, cooked into sauces Manganese 0.035 mg 2%
Phosphorus 11 mg 1%
and apple butter, or baked.[87 ] They are sometimes
Potassium 107 mg 4%
used as an ingredient in savory foods, such as sausage
Sodium 1 mg 0%
and stuffing.[88] Zinc 0.04 mg 0%

Several techniques are used to preserve apples and Other constituents Quantity
apple products. Traditional methods include drying and Water 85.56 g
making apple butter.[86] Juice and cider are produced
commercially; cider is a significant industry in regions Link to Full Nutrient Report of USDA
Database entry (https://fdc.nal.usda.gov/fdc-
such as the West of England and Normandy.[86] app.html#/food-details/1102644/nutrients)

A toffee apple (UK) or caramel apple (US) is a †

Percentages estimated using
confection made by coating an apple in hot toffee or US recommendations for adults, [84] except for
caramel candy respectively and allowing it to cool.[89][8] potassium, which is estimated based on expert
recommendation from the National Academies. [85]
Apples and honey are a ritual food pairing eaten during
the Jewish New Year of Rosh Hashanah.[90]

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.[86]

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 (called hard cider in North America),
ciderkin, and vinegar.[8] Through distillation, various alcoholic beverages can be produced, such as
applejack, Calvados, and apple brandy.[8][91 ]
Organic production
Organic apples are commonly produced in the United States.[92] Due to infestations by key insects and
diseases, organic production is difficult in Europe.[93] 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.[93] A
light coating of kaolin, which forms a physical barrier to some pests, also may help prevent apple sun
scalding.[48]

Non-browning apples
Apple skins and seeds contain polyphenols.[94] These are oxidised by the enzyme polyphenol oxidase,
which causes browning in sliced or bruised apples, by catalyzing the oxidation of phenolic compounds
to o-quinones, a browning factor.[95] 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.[96][97 ] The US Food and Drug Administration in 2015, and
Canadian Food Inspection Agency in 2017, determined that Arctic apples are as safe and nutritious as
conventional apples.[98][99]

Other products
Apple seed oil is obtained by pressing apple seeds for manufacturing cosmetics.[1 00]

In culture

Germanic paganism
In Norse mythology, the goddess Iðunn is portrayed in the Prose Edda
(written in the 13th century by Snorri Sturluson) as providing apples to
the gods that give them eternal youthfulness. The English scholar H. R.
Ellis Davidson links apples to religious practices in Germanic paganism,
from which Norse paganism developed. She points out that buckets of
apples were found in the Oseberg ship burial site in Norway, that fruit
and nuts (Iðunn having been described as being transformed into a nut in
Skáldskaparmál) have been found in the early graves of the Germanic
peoples in England and elsewhere on the continent of Europe, which may
have had a symbolic meaning, and that nuts are still a recognized symbol
of fertility in southwest England.[1 01 ]

Davidson notes a connection between apples and the Vanir, a tribe of gods "Brita as Iduna" (1901) by
associated with fertility in Norse mythology, citing an instance of eleven Carl Larsson

"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.[1 01 ]
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.[1 02]

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."[1 01 ]

Greek mythology
Apples appear in many religious traditions, including Greek and Roman
mythology where it has an ambiguous symbolism of discord, fertility, or
courtship.[1 03] 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.[1 04]

The Greek goddess of discord, Eris, became disgruntled after she was
excluded from the wedding of Peleus and Thetis.[1 05] 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, 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
Heracles with the apple of
Trojan War.[1 06][1 07 ] Hesperides

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:[1 08]

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),[1 04] 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.[1 09][1 1 0]

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".[1 1 1 ]

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.[3][1 1 2] 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
Píngānguǒ ("Peace apples") on sale
a way to wish them peace and safety.[1 1 2] in Beijing for Christmas Eve (2017)

Christian art
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.[1 1 3] The origin of the
popular identification with a fruit unknown in the Middle East in
biblical times is found in wordplay with the Latin words mālum (an
apple) and mălum (an evil), each of which is normally written
malum.[1 1 4] The tree of the forbidden fruit is called "the tree of
the knowledge of good and evil" in Genesis 2:17,[1 1 5] and the Latin
for "good and evil" is bonum et malum.[1 1 6]

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 Adam and Eve by Albrecht Dürer
sin itself. The larynx in the human throat has been called the (1507), showcasing the apple as a
"Adam's apple" because of a notion that it was caused by the symbol of sin
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.[1 1 3]

Proverb
The proverb, "An apple a day keeps the doctor away", addressing the supposed health benefits of the
fruit, has been traced to 19th-century Wales, where the original phrase was "Eat an apple on going to
bed, and you'll keep the doctor from earning his bread".[1 1 7 ] 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.[1 1 8]

See also
Apple chip
Applecrab, apple–crabapple hybrids for eating
Johnny Appleseed

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Further reading
Browning, Frank (1998). Apples (https://archive.org/details/apples00brow) (First ed.). New York:
North Point Press. ISBN 978-0-86547-537-3. LCCN 98027252 (https://lccn.loc.gov/98027252).
OCLC 39235786 (https://search.worldcat.org/oclc/39235786).
Hanson, Beth; Marinelli, Janet; Saphire, Sigrun Wolff; Tebbitt, Mark, eds. (2003). The Best Apples to
Buy and Grow (https://archive.org/details/bestapplestobuyg0000unse) (First ed.). Brooklyn, New
York: Brooklyn Botanic Garden. ISBN 978-1-889538-66-2. OCLC 60384060 (https://search.worldcat.
org/oclc/60384060).
Juniper, Barrie E.; Mabberley, David J. (2006). The Story of the Apple (https://archive.org/details/stor
yofapple0000juni) (First ed.). Portland, Oregon: Timber Press. ISBN 978-0-88192-784-9.
LCCN 2006011869 (https://lccn.loc.gov/2006011869). OCLC 67383484 (https://search.worldcat.org/
oclc/67383484).
Phillips, Michael (1998). The Apple Grower : A Guide for the Organic Orchardist (https://archive.org/d
etails/applegrowerguide0000phil) (First ed.). White River Junction, Vermont: Chelsea Green
Publishing. ISBN 978-1-890132-04-0. LCCN 98003631 (https://lccn.loc.gov/98003631).
OCLC 38731995 (https://search.worldcat.org/oclc/38731995).
Sanders, Rosie (2010). The Apple Book (Second ed.). London: Frances Lincoln Limited.
ISBN 9780711231412. OCLC 646397065 (https://search.worldcat.org/oclc/646397065).

External links
Media related to Apples at Wikimedia Commons

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