Apple

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, where its wild ancestor, Malus sieversii, is still
found. Apples have been grown for thousands of years in Eurasia before they were introduced to
North America by European colonists. Apples have cultural significance in many mythologies
(including Norse and Greek) and religions (such as Christianity in Europe).

Apples grown from seeds 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, eating raw, and cider 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 ancestor is æppel, is descended from the Proto-Germanic
noun *aplaz, descended in turn from Proto-Indo-European *h₂ébōl.[3] 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 expression appel of paradis, meaning a banana.[4]

Description

The apple is a deciduous tree, generally standing 2 to 4.5 metres (6 to 15 feet) tall in cultivation
and up to 15 m (49 ft) in the wild, though more typically 2 to 10 m (6.5 to 33 ft).[5][1] When
cultivated, the size, shape and branch density are determined by rootstock selection and
trimming method.[5] Apple trees may naturally have a rounded to erect crown with a dense
canopy of leaves.[6] The bark of the trunk is dark gray 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
Apple
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- 'Cripps Pink' cultivar

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-11⁄2 in) long.[6][1] Flowers of M. domestica

Blossoms are produced in spring simultaneously with Scientific classification

the budding of the leaves and are produced on spurs

Kingdom: Plantae
and some long shoots.[5] When the flower buds first
begin to open the petals are rose-pink and fade to Clade: Tracheophytes
white or light pink when fully open with each flower 3-
Clade: Angiosperms
to-4-centimeter (1-to-11⁄2-inch) in diameter.[1] The
five-petaled flowers are group in an inflorescence Clade: Eudicots
[8]
consisting of a cyme with 3–7 flowers. The central
flower of the inflorescence is called the "king bloom"; Clade: Rosids

it opens first and can develop a larger fruit.[6] Open

Order: Rosales
apple blossoms are damaged by even brief
exposures to temperatures −2 °C (28 °F) or less, Family: Rosaceae
although the overwintering wood and buds are hardy
Genus: Malus
down to −40 °C (−40 °F).[8]
Species: M. domestica

Binomial name

Malus domestica
(Suckow) Borkh.

Synonyms[1][2]

M. communis Desf., 1768

 M. pumila Mil.

M. frutescens Medik.

M. paradisiaca (L.) Medikus

M. sylvestris Mil.

Pyrus malus L.

Pyrus malus var. paradisiaca L.

Pyrus dioica Moench

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.[10]

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.[11]
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.[12]

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%.[13] The skin may be wholly or partly russeted, making it rough and brown. The skin is
covered in a protective layer of epicuticular wax.[14] 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.[13]

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.[15][16]

Taxonomy

The apple as a species has more than 100 alternative scientific names, or synonyms.[17] 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.[18] The General Committee of the IAPT decided in June 2017
to approve this change, officially conserving M. domestica.[19] 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.[17] 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]

Wild Malus sieversii apple in Kazakhstan

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 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.[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]

"Wild Apples"
by Henry David Thoreau
Read by Kevin S for LibriVox

1:01:36

Audio 01:01:35 (full text (https://archive.org/details/excursions1863thor/page/266) )

Problems playing this file? See media help.

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.[33] 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.[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

An apple tree in Germany

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 (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 An orchard mason bee on an

cultivar 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

Rome apples on sale at a farmer's market

in Newark, Delaware, United States (2011)

Commercially, apples can be stored for months in controlled atmosphere chambers. Apples are
commonly stored in chambers with lowered concentrations of oxygen to reduce respiration and
slow softening and other changes if the fruit is already fully ripe. The gas ethylene is used by
plants as a hormone which promotes ripening, decreasing the time an apple can be stored. For
storage longer than about six months the apples are picked earlier, before full ripeness, when
ethylene production by the fruit is low. However, in many varieties this increases their sensitivity
to carbon dioxide, which also must be controlled.[53]

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

Codling moth larva tunnelling inside an

apple

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.[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

From left to right: the 'Golden Delicious',

'SweeTango', 'Granny Smith', and 'Gala'
apples.

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 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.[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.[70]

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.[71] Released by the University of Minnesota in 1991, the
'Honeycrisp' has become famous for its crispness and juiciness, thereby commanding high
market prices.[71] Unusually for a popular cultivar, the 'Honeycrisp' is not directly related to
another popular apple cultivar but instead to two unsuccessful cultivars.[72] 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.[73] By the
2020s, about half of the new apple varieties entering the market in the United States and Canada
are 'Honeycrisp' progeny.[74] 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.[73]

Less common apple cultivars from an

orchard in Italy

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.[75] Most North Americans and Europeans favor crunchy, sweet, and subacid apples.[76]
Nevertheless, tart apples maintain a strong minority following.[77] In the United States today, the
most popular apple varieties are the 'Ambrosia', 'Honeycrisp', and 'Jazz', according Nielsen
data,[73] while in Canada, the 'Honeyscrisp', 'Ambrosia', and 'Gala' take the top spots.[75] Together,
these relatively newer varieties have overtaken once dominant cultivars like the 'McIntosh' and
the 'Red Delicious' in the North American market.[75][76] Extremely sweet apples with barely any
acid flavor are popular in Asia,[77] 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.[78] 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
Apple production
World production of apples in 2023 was 97 million tonnes, with China
producing 51% of the total (table).[79] Secondary producers were the 2023, millions of tonnes
[79]
United States, Turkey, and Poland.
China 49.6

United States 5.2

Toxicity
Turkey 4.6

Poland 3.9
Amygdalin India 2.9

Iran 2.2
Apple seeds contain small amounts of amygdalin, a sugar and cyanide
compound known as a cyanogenic glycoside. Ingesting small amounts World 97.3

of apple seeds causes no ill effects, but consumption of extremely Source: FAOSTAT of the United
Nations[79]
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.[80] The U.S. National Library of Medicine's
Hazardous Substances Data Bank records no cases of amygdalin poisoning from consuming
apple seeds.[81]

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.[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 allergies to other fruits, nuts, and vegetables. Reactions, which entail oral allergy
syndrome (OAS), generally involve itching and inflammation of the mouth and throat,[82] but in
rare cases can also include life-threatening anaphylaxis.[83] 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.[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

Apples, with skin (edible parts)

Nutritional value per 100 g (3.5 oz)

Energy 218 kJ (52 kcal)

Carbohydrates 13.81 g

Sugars 10.39

Dietary fiber 2.4 g

Fat 0.17 g

Protein 0.26 g

Vitamins and minerals [show]

Other constituents Quantity

Water 85.56 g

Link to Full Nutrient Report of USDA

Database entry (https://fdc.nal.usda.gov/fdc-
app.html#/food-details/1102644/nutrient
s)
†
Percentages estimated using US recommendations
for adults,[84] except for potassium, which is
estimated based on expert recommendation from the
National Academies.[85]

A raw apple is 86% water and 14% carbohydrates, with negligible content of fat and protein
(table). A reference serving of a raw apple with skin weighing 100 g (3.5 oz) 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

Machine for paring, coring, and slicing

apples, from Henry B. Scammell's 1897
handbook Cyclopedia of Valuable Receipts

Apples varieties can be grouped as cooking apples, eating apples, and cider apples, the last so
astringent as to be "almost inedible".[86] Apples are consumed as juice, raw in salads, baked in
pies, cooked into sauces and apple butter, or baked.[87] They are sometimes used as an
ingredient in savory foods, such as sausage and stuffing.[88]

Several techniques are used to preserve apples and apple products. Traditional methods include
drying and making apple butter.[86] Juice and cider are produced commercially; cider is a
significant industry in regions such as the West of England and Normandy.[86]

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.[89][8] 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.[100]

In culture

Germanic paganism

"Brita as Iduna" (1901) by Carl

Larsson

In Norse mythology, the goddess Iðunn is portrayed in the Prose Edda (written in the 13th century
by Snorri Sturluson) as providing apples to the gods that give them eternal youthfulness. 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, and that fruits and nuts have been found in the early graves of
the Germanic peoples 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.[101]

Davidson notes a connection between apples and the Vanir, a tribe of gods associated with
fertility in Norse mythology, citing an instance of eleven "golden apples" being given to woo the
beautiful Gerðr by Skírnir, who was acting as messenger for the major Vanir god Freyr in stanzas
19 and 20 of Skírnismál. Davidson also notes a further connection between fertility and apples in
Norse mythology in chapter 2 of the Völsunga saga: when the major goddess Frigg sends King
Rerir an apple after he prays to Odin for a child, Frigg's messenger (in the guise of a crow) drops
the apple in his lap as he sits atop a mound.[101] 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.[102]

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."[101]
Greek mythology

Heracles with the apple of

Hesperides

Apples appear in many religious traditions, including Greek and Roman mythology where it has
an ambiguous symbolism of discord, fertility, or courtship.[103] 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.[104]

The Greek goddess of discord, Eris, became disgruntled after she was excluded from the
wedding of Peleus and Thetis.[105] 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 Trojan
War.[106][107]

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:[108]

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),[104] 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.[109][110]

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".[111]

China

Píngānguǒ ("Peace apples") on sale in

Beijing for Christmas Eve (2017)

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][112] 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.[112]
Christian art

Adam and Eve by Albrecht Dürer (1507),

showcasing the apple as a symbol of sin

Though the forbidden fruit of Eden in the Book of Genesis is not identified, popular Christian
tradition has held that it was an apple that Eve coaxed Adam to share with her.[113] 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.[114] The tree of the forbidden fruit is called "the tree of the knowledge of good and
evil" in Genesis 2:17,[115] and the Latin for "good and evil" is bonum et malum.[116]

Renaissance painters may also have been influenced by the story of the golden apples in the
Garden of Hesperides. As a result, in the story of Adam and Eve, the apple became a symbol for
knowledge, immortality, temptation, the fall of man into sin, and sin itself. The larynx in the
human throat has been called the "Adam's apple" because of a notion that it was caused by the
forbidden fruit remaining in the throat of Adam. The apple as symbol of sexual seduction has
been used to imply human sexuality, possibly in an ironic vein.[113]

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".[117] 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.[118]
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/98027
252) . 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 (http
s://search.worldcat.org/oclc/60384060) .

Juniper, Barrie E.; Mabberley, David J. (2006). The Story of the Apple (https://archive.org/detail
s/storyofapple0000juni) (First ed.). Portland, Oregon: Timber Press. ISBN 978-0-88192-784-
9. LCCN 2006011869 (https://lccn.loc.gov/2006011869) . OCLC 67383484 (https://search.w
orldcat.org/oclc/67383484) .

Phillips, Michael (1998). The Apple Grower : A Guide for the Organic Orchardist (https://archive.o
rg/details/applegrowerguide0000phil) (First ed.). White River Junction, Vermont: Chelsea
Green Publishing. ISBN 978-1-890132-04-0. LCCN 98003631 (https://lccn.loc.gov/9800363
1) . 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

Apple at the Wikibooks Cookbook subproject