Apple

{{Short description|Edible fruit}} {{hatnote group| {{About|the fruit|the technology company|Apple Inc.|other uses}} {{Redirect|Apple tree}} }} {{pp-semi-indef}} {{pp-move}} {{Use dmy dates|date=September 2019}} {{Good article}} {{Speciesbox | name = Apple | image = Pink lady and cross section.jpg | image_caption = 'Cripps Pink' cultivar | image2 = Malus domestica a1.jpg{{!}}class=notpageimage  | image2_caption = Flowers of ''M. domestica'' | genus = Malus | species = domestica | authority = (Suckow) Borkh., 1803<ref name="gbif">{{cite web|url=https://www.gbif.org/species/105528904/|title=''Malus domestica'' (Suckow) Borkh., 1803|publisher=Global Biodiversity Information Facility|date=2025|access-date=14 July 2025}}</ref> | synonyms = * ''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 | synonyms_ref = <ref name="FNA" /><ref name="POWO" /> }}

An '''apple''' is the round, edible fruit of 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'' is derived from Old English {{lang|ang-Latn|æppel}}, meaning "fruit", not specifically the apple.<ref name="OnlineEtymDict"/> That in turn is descended from the Proto-Germanic noun {{lang|gem-x-proto|*aplaz}}, descended in turn from Proto-Indo-European {{lang|ine-x-proto|*h₂ébōl}}.<ref name="Lisa Lim">{{cite news |last1=Lim |first1=Lisa |date=6 July 2021 |title=Where the word 'apple' came from and why the forbidden fruit was unlucky to be linked with the fall of man |url=https://www.scmp.com/magazines/post-magazine/article/3139890/where-word-apple-came-and-why-forbidden-fruit-was-unlucky |url-status=live |department=Language Matters |work=South China Morning Post |language=en |location=Hong Kong, China |publisher=Alibaba Group |archive-url=https://web.archive.org/web/20230628112156/https://www.scmp.com/magazines/post-magazine/article/3139890/where-word-apple-came-and-why-forbidden-fruit-was-unlucky |archive-date=28 June 2023 |access-date=28 June 2023}}</ref> As late as the 17th century, the word also functioned as a generic term for all fruit including nuts; one example is the pineapple, the swollen accessory fruit outside of the cashew nut known as the "cashew apple",<ref name="mw">{{cite web |title=A pineapple is an apple (kind of). How did this tropical fruit get tied to the apple? |url=https://www.merriam-webster.com/wordplay/word-history-pineapple |date=2025 |website=Word History |publisher=Merriam-Webster |access-date=11 April 2026 }}</ref> another is a 14th-century Middle English expression {{lang|enm|appel of paradis}}, meaning a banana.<ref name="OnlineEtymDict">{{cite web |title=Origin and history of "apple" by Online Etymology Dictionary |url=https://www.etymonline.com/word/apple |website=Online Etymology Dictionary |access-date=12 July 2025 |archive-date=21 December 2019 |archive-url=https://web.archive.org/web/20191221020212/https://www.etymonline.com/word/apple |url-status=live }}</ref>

== Description ==

The apple tree is deciduous, generally standing from {{convert|6|to|15|ft|m|abbr=off|order=flip|round=0.5}} tall in cultivation and up to {{cvt|15|m|ft|round=0.5}} in the wild, though more typically {{cvt|2 to 10|m|ft|round=0.5}}.<ref name="UofGeorgia" /><ref name="FNA">{{cite web |last=Dickson |first=Elizabeth E. |title=''Malus domestica'' |url=http://floranorthamerica.org/Malus_domestica |website=Flora of North America |access-date=27 July 2024 |date=28 May 2021 |url-status=live |archive-url=https://web.archive.org/web/20240728060606/http://floranorthamerica.org/Malus_domestica |archive-date=28 July 2024}}</ref> When cultivated, the size, shape and branch density are determined by rootstock selection and trimming method.<ref name="UofGeorgia" /> Apple trees may naturally have a rounded to erect crown with a dense canopy of leaves.<ref name="NC Extension" /> The bark of the trunk is dark gray or gray-brown, but young branches are reddish or dark-brown with a smooth texture.<ref name="FNA" /><ref name="Heil et al." /> Young twigs are covered in fine downy hairs; they become hairless when older.<ref name="Heil et al.">{{cite book |last1=Heil |first1=Kenneth D. |last2=O'Kane, Jr. |first2=Steve L. |last3=Reeves |first3=Linda Mary |last4=Clifford |first4=Arnold |date=2013 |title=Flora of the Four Corners Region: Vascular Plants of the San Juan River Drainage, Arizona, Colorado, New Mexico, and Utah |url=https://archive.org/details/mobot31753003888887/page/n926 |edition=First |location=St. Louis, Missouri |publisher=Missouri Botanical Garden |page=909 |isbn=978-1-930723-84-9 |oclc=859541992 |access-date=27 July 2024}}</ref>

The buds are egg-shaped and dark red or purple in color; they range in size from 3 to 5mm, but are usually less than 4mm. The bud scales have very hairy edges. When emerging from the buds, the leaves are {{plantgloss|convolute}}, meaning that their edges overlap each other.<ref name="FNA" /> 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.<ref name="Heil et al." /><ref name="FNA" /> The edges have broadly-angled teeth, but do not have lobes. The top surface of the leaves are {{plantgloss|glabrescent}}, almost hairless, while the undersides are densely covered in fine hairs.<ref name="FNA" /> The leaves are attached alternately by short leaf stems {{cvt|1 to 3.5|cm||sp=us|adj=on|frac=2}} long.<ref name="NC Extension" /><ref name="FNA" />

Blossoms are produced in spring simultaneously with the budding of the leaves and are produced on spurs and some long shoots.<ref name="UofGeorgia" /> 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 {{convert|3|to|4|cm|abbr=off|sp=us|adj=on|frac=2}} in diameter.<ref name="FNA" /> The five-petaled flowers are group in an inflorescence consisting of a cyme with 3–7 flowers.<ref name="Lim 2012">{{cite book |last1=Lim |first1=Tong Kwee |date=2012 |chapter=''Malus'' x ''domestica'' |title=Edible Medicinal and Non-Medicinal Plants |language=en |volume=4, Fruit |edition=First |location=Dordrecht, the Netherlands |publisher=Springer |pages=414–415 |doi=10.1007/978-94-007-4053-2_49 |isbn=978-94-007-4053-2 |oclc=795503871}}</ref> The central flower of the inflorescence is called the "king bloom"; it opens first and can develop a larger fruit.<ref name="NC Extension">{{cite web |title=Apples - ''Malus domestica'' |url=https://plants.ces.ncsu.edu/plants/malus-domestica/common-name/apples/ |website=North Carolina Extension Gardener Plant Toolbox |publisher=North Carolina State University |access-date=31 July 2024 |archive-url=https://web.archive.org/web/20240531122432/https://plants.ces.ncsu.edu/plants/malus-domestica/common-name/apples/ |archive-date=31 May 2024 |url-status=live}}</ref> Open apple blossoms are damaged by even brief exposures to temperatures {{cvt|-2|C|F}} or less, although the overwintering wood and buds are hardy down to {{cvt|-40|C|F}}.<ref name="Lim 2012" />

{{gallery|mode=packed |Apple Blossom @ Manali.jpg|Apple blossoms |Koeh-108.jpg|Botanical illustration }}

=== Fruit ===

The fruit is a pome that matures in late summer or autumn.<ref name="FNA" /> 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.<ref>{{cite book |last1=Juniper |first1=Barrie E. |last2=Mabberley |first2=David J. |author1-link=Barrie Juniper |author2-link=David Mabberley |date=2006 |title=The Story of the Apple |url=https://archive.org/details/storyofapple0000juni |url-access=registration |edition=First |location=Portland, Oregon |publisher=Timber Press |page=27 |isbn=978-0-88192-784-9 |lccn=2006011869 |oclc=67383484 |access-date=1 August 2024}}</ref> The edible flesh is formed from the receptacle at the base of the flower.<ref>{{cite web |title=Fruit glossary |url=https://schoolgardening.rhs.org.uk/Resources/Info-Sheet/Fruit-Glossary |publisher=Royal Horticultural Society |access-date=7 August 2024 |url-status=live |archive-url=https://web.archive.org/web/20240807071550/https://schoolgardening.rhs.org.uk/Resources/Info-Sheet/Fruit-Glossary |archive-date=7 August 2024}}</ref>

{{gallery|mode=packed|align=center|width=400|height=250| File:Apple anatomy, flower and fruit compared.svg|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.<ref>{{cite book |last1=Burford |first1=Tom |author1-link=Tom Burford |date=2013 |title=Apples of North America: 192 Exceptional Varieties for Gardeners, Growers and Cooks |url=https://archive.org/details/applesofnorthame0000burf |url-access=limited |language=en |edition=First |location=Portland, Oregon |publisher=Timber Press |pages=22, 50, 55, 122, 123, 137, 141, 147, 159, 245, 246 |isbn=978-1-60469-249-5 |lccn=2012045130 |oclc=819860825}}</ref> The five sepals remain attached and stand out from the surface of the apple.<ref name="FNA"/>

The size of the fruit varies widely between cultivars, but generally has a diameter between {{cvt|2.5 and 12|cm|in|0}}.<ref name="Heil et al."/> The shape is quite variable and may be nearly round, elongated, conical, or short and wide.<ref>{{cite web |title=Shape |url=https://agresearch.montana.edu/warc/guides/apples/heritage_orchard_management_guide/heritage-fruit-id-guide/apple_id_characteristics/appleid_shape.html |website=Western Agricultural Research Center |publisher=Montana State University |access-date=30 July 2024 |url-status=live |archive-url=https://web.archive.org/web/20240423183929/https://agresearch.montana.edu/warc/guides/apples/heritage_orchard_management_guide/heritage-fruit-id-guide/apple_id_characteristics/appleid_shape.html |archive-date=23 April 2024}}</ref>

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%.<ref name="Janick">{{cite book |last1=Janick |first1=Jules |last2=Cummins |first2=James N. |last3=Brown |first3=Susan K. |last4=Hemmat |first4=Minou |date=1996 |chapter=Chapter 1: Apples |chapter-url=http://www.hort.purdue.edu/newcrop/pri/chapter.pdf |title=Fruit Breeding |url-status=live |volume=I: Tree and Tropical Fruits |location=New York |publisher=John Wiley & Sons |pages=9, 48 |isbn=978-0-471-31014-3 |lccn=95016407 |oclc=1302621533 |archive-url=https://web.archive.org/web/20130719085631/http://www.hort.purdue.edu/newcrop/pri/chapter.pdf |archive-date=19 July 2013 |access-date=30 August 2024}}</ref> The skin may be wholly or partly russeted, making it rough and brown. The skin is covered in a protective layer of epicuticular wax.<ref>{{Cite web |url=http://postharvest.tfrec.wsu.edu/pages/N2I2A |title=Natural Waxes on Fruits |last1=Kolattukudy |first1=P. E. |date=2013 |orig-date=May 1984 |website=Postharvest Information Network |publisher=Washington State University Tree Fruit Research & Extension Center |language=en |access-date=14 June 2013 |archive-url=https://web.archive.org/web/20130524130738/http://postharvest.tfrec.wsu.edu/pages/N2I2A |archive-date=24 May 2013}}</ref> The skin may also be marked with scattered dots.<ref name="FNA" /> The flesh is generally pale yellowish-white, though it can be pink, yellow or green.<ref name="Janick" />

{{gallery|align=center|mode=nolines|width=150|title=Apples can have any amount of overcolor, a darker tint over a pale groundcolor.| File:Yellow Transparent (cropped).jpg|0% overcolor| File:Lobo (apple) (cropped).jpg|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.<ref>{{cite journal |last1=Flath |first1=R. A. |last2=Black |first2=D. R. |last3=Forrey |first3=R. R. |last4=McDonald |first4=G. M. |last5=Mon |first5=T. R. |last6=Teranishi |first6=R. |title=Volatiles in Gravenstein Apple Essence Identified by GC-Mass Spectrometry |journal=Journal of Chromatographic Science |date=1 August 1969 |volume=7 |issue=8 |page=508 |doi=10.1093/CHROMSCI/7.8.508}}</ref><ref>{{cite journal |last1=Flath |first1=Robert A. |last2=Black |first2=Dale Robert. |last3=Guadagni |first3=Dante G. |last4=McFadden |first4=William H. |last5=Schultz |first5=Thomas H. |title=Identification and organoleptic evaluation of compounds in Delicious apple essence |journal=Journal of Agricultural and Food Chemistry |date=January 1967 |volume=15 |issue=1 |page=29 |doi=10.1021/jf60149a032|bibcode=1967JAFC...15...29F }}</ref>

== Taxonomy == The apple as a species has more than 100 alternative scientific names, or synonyms.<ref name="Proposal to conserve">{{cite journal |last1=Qian |first1=Guan-Ze |last2=Liu |first2=Lian-Fen |last3=Tang |first3=Geng-Guo |title=(1933) Proposal to conserve the name ''Malus domestica'' against ''M. pumila'', ''M. communis'', ''M. frutescens'', and ''Pyrus dioica'' ( Rosaceae ) |journal=Taxon |date=April 2010 |volume=59 |issue=2 |pages=650–652 |doi=10.1002/tax.592038 |bibcode=2010Taxon..59..650Q |language=en}}</ref> 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.<ref>{{cite journal |url=https://apirs.plants.ifas.ufl.edu/site/assets/files/375067/375067.pdf |doi=10.12705/662.17 |title=Report of the Nomenclature Committee for Vascular Plants: 69 |date=2017 |last1=Applequist |first1=Wendy L. |journal=Taxon |volume=66 |issue=2 |pages=500–513 |bibcode=2017Taxon..66..500A |url-status=live |archive-url=https://web.archive.org/web/20240507042204/https://apirs.plants.ifas.ufl.edu/site/assets/files/375067/375067.pdf |archive-date=7 May 2024}}</ref> The General Committee of the IAPT decided in June 2017 to approve this change, officially conserving ''M. domestica''.<ref>{{cite journal |last1=Wilson |first1=Karen L. |title=Report of the General Committee: 18 |journal=Taxon |date=June 2017 |volume=66 |issue=3 |page=742 |doi=10.12705/663.15 |doi-access=free |bibcode=2017Taxon..66..742W }}</ref> Nevertheless, some works published after 2017 still use ''M. pumila'' as the correct name, under an alternate taxonomy.<ref name="POWO">{{cite POWO |id=726282-1 |title=''Malus domestica'' (Suckow) Borkh. |access-date=31 July 2024}}</ref>

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, the term 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.<ref name="Proposal to conserve" /> The earliest use of var. ''domestica'' for the apple was by Georg Adolf Suckow in 1786.<ref name="POWO" />

=== Genome === {{further |Apple 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'.<ref name="Velasco et al.">{{cite journal |year=2010 |title=The genome of the domesticated apple (''Malus'' × ''domestica'' Borkh.) |journal=Nature Genetics |doi=10.1038/ng.654 |display-authors=5 |last1=Velasco |first1=Riccardo |last2=Zharkikh |first2=Andrey |last3=Affourtit |first3=Jason |last4=Dhingra |first4=Amit |last5=Cestaro |first5=Alessandro |last6=Kalyanaraman |first6=Ananth |last7=Fontana |first7=Paolo |last8=Bhatnagar |first8=Satish K. |last9=Troggio |first9=Michela |last10=Pruss |first10=Dmitry |last11=Salvi |first11=Silvio |last12=Pindo |first12=Massimo |last13=Baldi |first13=Paolo |last14=Castelletti |first14=Sara |last15=Cavaiuolo |first15=Marina |last16=Coppola |first16=Giuseppina |last17=Costa |first17=Fabrizio |last18=Cova |first18=Valentina |last19=Dal Ri |first19=Antonio |last20=Goremykin |first20=Vadim |last21=Komjanc |first21=Matteo |last22=Longhi |first22=Sara |last23=Magnago |first23=Pierluigi |last24=Malacarne |first24=Giulia |last25=Malnoy |first25=Mickael |last26=Micheletti |first26=Diego |last27=Moretto |first27=Marco |last28=Perazzolli |first28=Michele |last29=Si-Ammour |first29=Azeddine |last30=Vezzulli |first30=Silvia |volume=42 |issue=10 |pages=833–839 |pmid=20802477 |s2cid=14854514 |doi-access=free}}</ref> However, this first whole genome sequence contained several errors,<ref>{{cite journal |year=2016 |title=A high-density, multi-parental SNP genetic map on apple validates a new mapping approach for outcrossing species |doi=10.1038/hortres.2016.57 |last1=Di Pierro |first1=Erica A. |last2=Gianfranceschi |first2=Luca |last3=Di Guardo |first3=Mario |last4=Koehorst-Van Putten |first4=Herma J.J. |last5=Kruisselbrink |first5=Johannes W. |last6=Longhi |first6=Sara |last7=Troggio |first7=Michela |last8=Bianco |first8=Luca |last9=Muranty |first9=Hélène |last10=Pagliarani |first10=Giulia |last11=Tartarini |first11=Stefano |last12=Letschka |first12=Thomas |last13=Lozano Luis |first13=Lidia |last14=Garkava-Gustavsson |first14=Larisa |last15=Micheletti |first15=Diego |last16=Bink |first16=Marco C.A.M. |last17=Voorrips |first17=Roeland E. |last18=Aziz |first18=Ebrahimi |last19=Velasco |first19=Riccardo |last20=Laurens |first20=François |last21=Van De Weg |first21=W. Eric |display-authors=5 |journal=Horticulture Research |volume=3 |issue=1 |page=16057 |pmid=27917289 |pmc=5120355 |bibcode=2016HorR....316057D }}</ref> 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.<ref name="Daccord et al.">{{cite journal |year=2017 |title=High-quality de novo assembly of the apple genome and methylome dynamics of early fruit development |publisher=Nature Communications |doi=10.1038/ng.3886 |last1=Daccord |first1=Nicolas |last2=Celton |first2=Jean-Marc |last3=Linsmith |first3=Gareth |last4=Becker |first4=Claude |last5=Choisne |first5=Nathalie |last6=Schijlen |first6=Elio |last7=Van De Geest |first7=Henri |last8=Bianco |first8=Luca |last9=Micheletti |first9=Diego |last10=Velasco |first10=Riccardo |last11=Di Pierro |first11=Erica Adele |last12=Gouzy |first12=Jérôme | last13=Rees |first13=D Jasper G. |last14=Guérif |first14=Philippe |last15=Muranty |first15=Hélène |last16=Durel |first16=Charles-Eric |last17=Laurens |first17=François |last18=Lespinasse |first18=Yves |last19=Gaillard |first19=Sylvain |last20=Aubourg |first20=Sébastien |last21=Quesneville |first21=Hadi |last22=Weigel |first22=Detlef |last23=Van De Weg |first23=Eric |last24=Troggio |first24=Michela |last25=Bucher |first25=Etienne |display-authors=3 |journal=Nature Genetics |volume=49 |issue=7 |pages=1099–1106 |pmid=28581499 |s2cid=24690391 |doi-access=free |hdl=10449/42064 |hdl-access=free}}</ref><ref name="Zhang et al.">{{cite journal |year=2019 |title=A high-quality apple genome assembly reveals the association of a retrotransposon and red fruit colour |publisher=Nature Genetics |doi=10.1038/s41467-019-09518-x |display-authors=5 |last1=Zhang |first1=Liyi |last2=Hu |first2=Jiang |last3=Han |first3=Xiaolei |last4=Li |first4=Jingjing |last5=Gao |first5=Yuan |last6=Richards |first6=Christopher M. |last7=Zhang |first7=Caixia |last8=Tian | first8=Yi |last9=Liu |first9=Guiming |last10=Gul |first10=Hera |last11=Wang |first11=Dajiang |last12=Tian |first12=Yu |last13=Yang |first13=Chuanxin |last14=Meng |first14=Minghui |last15=Yuan |first15=Gaopeng |last16=Kang |first16=Guodong |last17=Wu |first17=Yonglong |last18=Wang |first18=Kun |last19=Zhang |first19=Hengtao |last20=Wang |first20=Depeng |last21=Cong |first21=Peihua |journal=Nature Communications |volume=10 |issue=1 |page=1494 |pmid=30940818 |pmc=6445120 |bibcode=2019NatCo..10.1494Z}}</ref>

The first whole genome assembly was estimated to contain around 57,000 genes,<ref name="Velasco et al."/> though the more recent genome sequences support estimates between 42,000 and 44,700 protein-coding genes.<ref name="Daccord et al."/><ref name="Zhang et al."/> 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.<ref name="Duan 2017"/>

== Cultivation ==

=== History ===

thumb|center|upright=2.5|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.<ref name="Duan 2017"/>

[[File:95apple.jpeg|thumb|alt=color photograph of a hand holding a red apple|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.<ref name="Richards Volk 2009">{{cite journal |last1=Richards |first1=Christopher M. |display-authors=5 |author2=Volk, Gayle M. |author3=Reilley, Ann A. |author4=Henk, Adam D. |author5=Lockwood, Dale R. |author6=Reeves, Patrick A. |author7=Forsline, Philip L. |title=Genetic diversity and population structure in ''Malus sieversii'', a wild progenitor species of domesticated apple |journal=Tree Genetics & Genomes |volume=5 |issue=2 |pages=339–347 |year=2009 |doi=10.1007/s11295-008-0190-9 |s2cid=19847067}}</ref> 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.<ref name="UofGeorgia"/><ref>{{cite journal |last1=Lauri |first1=Pierre-éric |last2=Maguylo |first2=Karen |last3=Trottier |first3=Catherine |title=Architecture and size relations: an essay on the apple (Malus × domestica, Rosaceae) tree |journal=American Journal of Botany |date=March 2006 |volume=93 |issue=3 |pages=357–368 |doi=10.3732/ajb.93.3.357 |pmid=21646196 |bibcode=2006AmJB...93..357L |url=https://bsapubs.onlinelibrary.wiley.com/doi/full/10.3732/ajb.93.3.357 |access-date=27 July 2024 |archive-date=20 April 2019 |archive-url=https://web.archive.org/web/20190420154707/https://bsapubs.onlinelibrary.wiley.com/doi/full/10.3732/ajb.93.3.357 |url-status=live |url-access=subscription }}</ref> 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.<ref name="Cornille 2012">{{cite journal |last1=Cornille |first1=Amandine |title=New Insight into the History of Domesticated Apple: Secondary Contribution of the European Wild Apple to the Genome of Cultivated Varieties|journal=PLOS Genetics |doi=10.1371/journal.pgen.1002703 |year=2012 |editor1-last=Mauricio |editor1-first=Rodney |last2=Gladieux |first2=Pierre |last3=Smulders |first3=Marinus J. M. |last4=Roldán-Ruiz |first4=Isabel |last5=Laurens |first5=François |last6=Le Cam |first6=Bruno |last7=Nersesyan |first7=Anush |last8=Clavel |first8=Joanne |last9=Olonova |first9=Marina |volume=8 |issue=5 |article-number=e1002703 |pmid=22589740 |pmc=3349737 |display-authors=5 |doi-access=free }}</ref><ref>{{cite web |last=Kean |first=Sam |title=ScienceShot: The Secret History of the Domesticated Apple |date=17 May 2012 |url=https://www.science.org/content/article/scienceshot-secret-history-domesticated-apple |url-status=live |archive-url=https://web.archive.org/web/20160611184017/http://www.sciencemag.org/news/2012/05/scienceshot-secret-history-domesticated-apple |archive-date=11 June 2016}}</ref><ref>{{cite journal |last1=Coart |first1=E. |last2=Van Glabeke |first2=S. |last3=De Loose |first3=M. |last4=Larsen |first4=A.S. |last5=Roldán-Ruiz |first5=I. |year=2006 |title=Chloroplast diversity in the genus ''Malus'': new insights into the relationship between the European wild apple (''Malus sylvestris'' (L.) Mill.) and the domesticated apple (''Malus domestica'' Borkh.) |journal=Mol. Ecol. |volume=15 |issue=8 |pages=2171–2182 |doi=10.1111/j.1365-294x.2006.02924.x |pmid=16780433|bibcode=2006MolEc..15.2171C |s2cid=31481730 }}</ref>

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 hybridisation 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.<ref name="Duan 2017">{{cite journal |title=Genome re-sequencing reveals the history of apple and supports a two-stage model for fruit enlargement |doi=10.1038/s41467-017-00336-7|pmid=28811498|pmc=5557836 |journal=Nature Communications |year=2017 |last1=Duan |first1=Naibin |last2=Bai |first2=Yang |last3=Sun |first3=Honghe |last4=Wang |first4=Nan |last5=Ma |first5=Yumin |last6=Li |first6=Mingjun |last7=Wang |first7=Xin |last8=Jiao|first8=Chen |last9=Legall |first9=Noah |last10=Mao |first10=Linyong |last11=Wan |first11=Sibao |last12=Wang |first12=Kun |last13=He |first13=Tianming |last14=Feng |first14=Shouqian |last15=Zhang |first15=Zongying |last16=Mao |first16=Zhiquan |last17=Shen |first17=Xiang |last18=Chen |first18=Xiaoliu |last19=Jiang|first19=Yuanmao |last20=Wu |first20=Shujing |last21=Yin |first21=Chengmiao |last22=Ge |first22=Shunfeng |last23=Yang|first23=Long |last24=Jiang |first24=Shenghui |last25=Xu |first25=Haifeng |last26=Liu |first26=Jingxuan |last27=Wang |first27=Deyun |last28=Qu |first28=Changzhi |last29=Wang |first29=Yicheng |last30=Zuo |first30=Weifang |display-authors=5 |volume=8 |issue=1 |page=249 |bibcode=2017NatCo...8..249D}}</ref>

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.<ref name="Duan 2017"/>

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.<ref name="Duan 2017"/>

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.<ref>{{cite book |last1=Rottoli |first1=Mauro |last2=Pessina |first2=Andrea |editor-last1=Colledge |editor-first1=Sue |editor-last2=Conolly |editor-first2=James |date=2007 |chapter=Chapter 9: Neolithic agriculture in Italy: an update of archaeobotanical data with particular emphasis on northern settlements |title=The Origins and Spread of Domestic Plants in Southwest Asia and Europe |url=https://archive.org/details/originsspreadofd0000unse |url-access=limited |language=en |edition=First |location=Walnut Creek, California |publisher=Left Coast Press for the University College London Institute of Archaeology Publications |pages=142–143 |isbn=978-1-59874-988-5 |oclc=84838157}}</ref> 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.<ref name="Ancient DNA">{{cite journal |last1=Schlumbaum |first1=Angela |last2=van Glabeke |first2=Sabine |last3=Roldan-Ruiz |first3=Isabel |title=Towards the onset of fruit tree growing north of the Alps: Ancient DNA from waterlogged apple (''Malus'' sp.) seed fragments |journal=Annals of Anatomy - Anatomischer Anzeiger |date=January 2012 |volume=194 |issue=1 |pages=157–162 |doi=10.1016/j.aanat.2011.03.004 |pmid=21501956 |language=en}}</ref>

There is indirect evidence of apple cultivation in the third millennium BCE in the Middle East.<ref name="Ancient DNA"/> There is direct evidence, apple cores, dated to the 10th century BCE from a Judean site between the Sinai and Negev. <ref>{{cite book |last1=Sauer |first1=Jonathan D. |author-link1=Jonathan Deininger Sauer |date=1993 |title=Historical Geography of Crop Plants: A Select Roster |language=en |edition=First |location=Boca Raton, Florida |publisher=CRC Press |pages=109–113 |isbn=978-0-8493-8901-6 |lccn=92045590 |oclc=27224696}}</ref> There was substantial apple production in European classical antiquity, and grafting was certainly known then.<ref name="Ancient DNA"/> 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.<ref name="Ancient DNA"/>

{{listen | filename = Wild Apples by Henry David Thoreau - read by Kevin S for LibriVox's Short Nonfiction Collection Vol. 065 (2019).ogg | title = "Wild Apples" by Henry David Thoreau Read by Kevin S for LibriVox | description = Audio 01:01:35 ([https://archive.org/details/excursions1863thor/page/266 full text]) | pos = right | type = speech | image = 60px }}

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.<ref>{{cite book |last1=Plinius |first1=Gaius Secundus |author1-link=Pliny the Elder |translator-last1=Bostock |translator-first1=John |translator-link1=John Bostock (physician) |translator-last2=Riley |translator-first2=Henry T. |translator-link2=Henry Thomas Riley |title=The Natural History of Pliny |date=1855 |publisher=Henry G. Bohn |location=London |page=303 |url=https://archive.org/details/naturalhistoryof03plin/page/303 |access-date=3 August 2024 |volume=III}}</ref> These methods extend the shelf life of fresh apples, but refrigeration is still required. Even sturdy winter varieties only keep well until December in cool climates.<ref>{{cite book |last1=Martin |first1=Alice A. |date=1976 |title=All About Apples |url=https://archive.org/details/allaboutapples0000mart/page/64 |language=en |edition=First |location=Boston, Massachusetts |publisher=Houghton Mifflin Company |pages=64–65 |isbn=978-0-395-20724-6 |oclc=1733691 |access-date=3 August 2024}}</ref> For longer storage medieval Europeans strung up cored and peeled apples to dry, either whole or sliced into rings.<ref>{{cite book |last=Adamson |first=Melitta Weiss |date=2004 |title=Food in Medieval Times |url=https://archive.org/details/foodinmedievalti0000adam |url-access=limited |location=Westport, Connecticut |publisher=Greenwood Press |pages=19–20 |isbn=978-0-313-32147-4 |lccn=2004014054 |oclc=55738647}}</ref>

Of the many Old World plants that the Spanish introduced to Chiloé Archipelago in the 16th century, apple trees became particularly well adapted.<ref name=Torrejonetal2004>{{cite journal |last1=Torrejón |first1=Fernando |last2=Cisternas |first2=Marco |last3=Araneda |first3=Alberto |date=2004 |title=Efectos ambientales de la colonización española desde el río Maullín al archipiélago de Chiloé, sur de Chile |trans-title=Environmental effects of the spanish colonization from de Maullín river to the Chiloé archipelago, southern Chile |journal=Revista Chilena de Historia Natural |volume=77 |issue=4 |pages=661–677 |doi=10.4067/s0716-078x2004000400009 |language=es |doi-access=free |hdl=10533/175736 |hdl-access=free }}</ref> Apples were introduced to North America by colonists in the 17th century,<ref name="UofGeorgia" /> and the first named apple cultivar was introduced in Boston by Reverend William Blaxton in 1640.<ref>{{cite book |last1=Smith |first1=Archibald William |date=1963 |title=A Gardener's Book of Plant Names: A Handbook of the Meaning and Origins of Plant Names |url=https://archive.org/details/gardenersbookofp00smit/page/40 |url-access=registration |language=en |edition=First |location=New York |publisher=Harper & Row |page=40 |lccn=62009906 |oclc=710612 |access-date=10 August 2024}}</ref> The only apples native to North America are crab apples.<ref name="Harrowsmith">{{cite book |last1=Poole |first1=Mike |editor1-last=Lawrence |editor1-first=James |date=1980 |chapter=Heirloom Apples |chapter-url=https://archive.org/details/harrowsmithreade0000unse_k4b7/page/122 |chapter-url-access=registration |title=The Harrowsmith Reader Volume II |language=en |location=Camden East, Ontario |publisher=Camden House Publishing |page=122 |isbn=978-0-920656-11-2 |oclc=1336124440 |access-date=10 August 2024}}</ref>

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.<ref name="Harrowsmith" /> 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.<ref name="UofGeorgia"/>

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.<ref>{{cite book |last1=Van Valen |first1=James M. |date=1900 |title=History of Bergen County, New Jersey |url=https://archive.org/details/historybergenco00valegoog/page/33 |language=en |location=New York |publisher=New Jersey Publishing and Engraving Company |pages=33–34 |oclc=25697876 |access-date=9 August 2024}}</ref><ref>{{cite book |last1=Brox |first1=Jane |author1-link=Jane Brox |date=1999 |title=Five Thousand Days Like This One |url=https://archive.org/details/fivethousanddays00brox/page/150 |url-access=registration |language=en |edition=First |location=Boston, Massachusetts |publisher=Beacon Press |pages=150–151 |isbn=978-0-8070-2106-4 |lccn=98035051 |oclc=39605684 |access-date=9 August 2024}}</ref> 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.<ref>{{cite web |last1=Cohen |first1=Rachel D. |title=Thanks To Science, You Can Eat An Apple Every Day |url=https://www.npr.org/sections/thesalt/2018/11/26/668256349/thanks-to-science-you-can-eat-an-apple-every-day |website=The Salt |publisher=NPR |access-date=1 August 2024 |archive-url=https://web.archive.org/web/20240618003023/https://www.npr.org/sections/thesalt/2018/11/26/668256349/thanks-to-science-you-can-eat-an-apple-every-day |archive-date=18 June 2024 |date=26 November 2018 |url-status=live}}</ref>

=== Breeding === {{See also|Fruit tree propagation|Malling series}}

thumb|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 do not "breed true", instead they are "extreme heterozygotes", i.e. rather than resembling their parents, seedlings are all different from each other and from their parents.<ref>{{cite web |title=The Heirloom Apple Orchard |url=https://blogs.cornell.edu/jentsch/the-heirloom-orchard/ |website=The Jentsch Lab |publisher=Cornell University |access-date=9 August 2024 |archive-url=https://web.archive.org/web/20240730203237/https://blogs.cornell.edu/jentsch/the-heirloom-orchard/ |archive-date=30 July 2024 |url-status=live}}</ref> 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.<ref>{{cite conference |last=Ranney |first=Thomas G. |title=Polyploidy: From Evolution to Landscape Plant Improvement |url=http://www.ces.ncsu.edu/fletcher/programs/nursery/metria/metria11/ranney/index.html |conference=11th Metropolitan Tree Improvement Alliance Conference held in Gresham, Oregon, August 23–24, 2000 |publisher=METRIA |archive-url=https://web.archive.org/web/20100723170402/http://www.ces.ncsu.edu/fletcher/programs/nursery/metria/metria11/ranney/index.html |archive-date=23 July 2010 |access-date=7 November 2010 |conference-url=https://www.ces.ncsu.edu/fletcher/programs/nursery/metria/metria11/index.html |book-title=Proceedings of the 11th Metropolitan Tree Improvement Alliance (METRIA) Conference |website=METRIA (NCSU.edu) }}</ref>

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 {{convert|10|ft|m|order=flip|abbr=on|disp=or}} high at maturity), which bear fruit many years earlier in their life cycle than full size trees, and are easier to harvest.<ref>{{cite web |last1=Lord |first1=William G. |last2=Ouellette |first2=Amy |date=February 2010 |title=Dwarf Rootstocks for Apple Trees in the Home Garden |url=http://extension.unh.edu/resources/files/resource000583_rep605.pdf |archive-url=https://web.archive.org/web/20130930151829/http://extension.unh.edu/resources/files/Resource000583_Rep605.pdf |archive-date=30 September 2013 |access-date=1 September 2013 |publisher=University of New Hampshire}}</ref>

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.<ref>{{cite journal |last1=Fallahi |first1=Esmaeil |last2=Colt |first2=W. Michael |last3=Fallahi |first3=Bahar |last4=Chun |first4=Ik-Jo |title=The Importance of Apple Rootstocks on Tree Growth, Yield, Fruit Quality, Leaf Nutrition, and Photosynthesis with an Emphasis on 'Fuji' |journal=HortTechnology |date=January 2002 |volume=12 |issue=1 |pages=38–44 |doi=10.21273/HORTTECH.12.1.38 |url=https://journals.ashs.org/horttech/view/journals/horttech/12/1/article-p38.xml |access-date=9 August 2024 |url-status=live |archive-url=https://web.archive.org/web/20140211173706/http://horttech.ashspublications.org/content/12/1/38.full.pdf |archive-date=11 February 2014 |language=en}}</ref> 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.<ref>{{cite web |last1=Parker |first1=M.L. |title=Apple Rootstocks and Tree Spacing |url=http://www.ces.ncsu.edu/hil/hil-300-a.html |website=North Carolina Cooperative Extension Service |access-date=1 September 2013 |archive-url=https://web.archive.org/web/20130911051127/http://www.ces.ncsu.edu/hil/hil-300-a.html |archive-date=11 September 2013 |language=en |date=September 1993 }}</ref>

Most new apple cultivars originate as seedlings, which either arise by chance or are bred by deliberately crossing cultivars with promising characteristics.<ref>{{cite book |last1=Ferree |first1=David Curtis |last2=Warrington |first2=Ian J. |title=Apples: Botany, Production, and Uses |date=2003 |publisher=Centre for Agriculture and Bioscience International |location=New York |isbn=978-0-85199-592-2 |pages=33–35 |language=en |oclc=133167834}}</ref> 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.<ref name="Polomski & Reighard">{{cite web |last1=Polomski |first1=Bob |last2=Reighard |first2=Greg |title=Apple HGIC 1350 |url=http://hgic.clemson.edu/factsheets/HGIC1350.htm |website=Home & Garden Information Center |publisher=Clemson University |access-date=22 January 2008 |archive-url=https://web.archive.org/web/20080228131352/http://hgic.clemson.edu/factsheets/HGIC1350.htm |archive-date=28 February 2008 }}</ref>

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.<ref>{{cite journal |last1=Barahona |first1=M. |title=Adaptation of Apple Varieties in Ecuador |journal=Acta Horticulturae |date=1992 |issue=310 |pages=135–142 |doi=10.17660/ActaHortic.1992.310.17}}</ref>

=== Pollination ===

<gallery mode="packed" heights="200"> File:Apple tree blossom.JPG|An apple blossom from an old Ayrshire cultivar File:Orchmason.jpg|An orchard mason bee on an apple bloom </gallery>

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.<ref name="Polomski & Reighard" /><ref>{{cite thesis |last=Adamson |first=Nancy Lee |date=2011 |title=An Assessment of Non-Apis Bees as Fruit and Vegetable Crop Pollinators in Southwest Virginia |url=http://www.step-project.net/NPDOCS/Adamson_NL_D_2011.pdf |degree=Doctor of Philosophy in Entomology |publisher=Virginia Polytechnic Institute and State University |docket= |oclc= |access-date=15 October 2015 |archive-url=https://web.archive.org/web/20151120230411/http://www.step-project.net/NPDOCS/Adamson_NL_D_2011.pdf |archive-date=20 November 2015 |url-status=live }}</ref>

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:<ref name="Powell 1986"/>

* 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).<ref name="Powell 1986">{{cite journal |last1=Powell |first1=L.E. |date=1986 |title=The Chilling Requirement in Apple and Its Role in Regulating Time of Flowering in Spring in Cold-Winter Climate |journal=Acta Horticulturae |location=Wageningen, Netherlands |publisher=International Society for Horticultural Science |issue=179 |pages=129–140 |doi=10.17660/ActaHortic.1986.179.10 |isbn=978-90-6605-182-9}}</ref>

=== Maturation and harvest ===

<gallery mode="packed" heights="200"> 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) </gallery>

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 {{convert|40|-|200|kg|lb|-1|abbr=on}} 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 {{convert|10|-|80|kg|lb|-1|abbr=on}} of fruit per year.<ref name="Polomski & Reighard" />

Some farms with apple orchards open them to the public so consumers can pick their own apples.<ref>{{cite web |last=Romano |first=Andrea |title=20 Best Places to Go Apple Picking in the United States |url=https://www.travelandleisure.com/trip-ideas/fall-vacations/best-orchards-for-apple-picking |website=Travel + Leisure |access-date=2 August 2024 |archive-url=https://web.archive.org/web/20240421140207/https://www.travelandleisure.com/trip-ideas/fall-vacations/best-orchards-for-apple-picking |archive-date=21 April 2024 |date=10 September 2023 |url-status=dead}}</ref>

Crops ripen at different times of the year according to the cultivar. Cultivars that yield their crop in the summer include 'Sweet Bough' and 'Duchess'; fall producers include 'Blenheim'; winter producers include 'King', 'Swayzie', and 'Tolman Sweet'.<ref name="Harrowsmith"/>

=== Storage === [[File:Rome Apples, Newark Delaware Farmer's Market.jpg|thumb|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.<ref>{{cite web |last1=Graziano |first1=Jack |last2=Farcuh |first2=Macarena |title=Controlled Atmosphere Storage of Apples |url=https://extension.umd.edu/resource/controlled-atmosphere-storage-apples |website=University of Maryland Extension |access-date=2 August 2024 |archive-url=https://web.archive.org/web/20230324131954/https://extension.umd.edu/resource/controlled-atmosphere-storage-apples |archive-date=24 March 2023 |date=10 September 2021 |url-status=live}}</ref>

For home storage, most cultivars 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 {{cvt|4 to 0|C|F}}.<ref>{{cite web |title=FoodKeeper App |url=https://www.foodsafety.gov/keep-food-safe/foodkeeper-app |website=FoodSafety.gov |publisher=United States Department of Health and Human Services |access-date=17 September 2024 |language=en |date=26 April 2019}}</ref><ref>{{cite web |title=4 Steps to Food Safety |url=https://www.foodsafety.gov/keep-food-safe/4-steps-to-food-safety |website=FoodSafety.gov |publisher=United States Department of Health and Human Services |access-date=17 September 2024 |language=en |date=12 April 2019}}</ref> Some varieties of apples (e.g. 'Granny Smith' and 'Fuji') have more than three times the storage life of others.<ref>{{cite web |access-date=25 May 2007 |url=http://www.csiro.au/en/Research/Health/Food-safety/Refrigerating-foods |title=Refrigerated storage of perishable foods |date=26 February 2015 |publisher=CSIRO |archive-url=https://web.archive.org/web/20150315033439/http://www.csiro.au/en/Research/Health/Food-safety/Refrigerating-foods |archive-date=15 March 2015 }}</ref>

Non-organic apples may be sprayed with a substance 1-methylcyclopropene blocking the apples' ethylene receptors, temporarily preventing them from ripening.<ref>{{cite news |last1=Karp |first1=David |date=25 October 2006 |title=Puff the Magic Preservative: Lasting Crunch, but Less Scent |work=The New York Times |url=https://www.nytimes.com/2006/10/25/dining/25fruit.html |url-status=live |access-date=26 July 2017 |archive-url=https://web.archive.org/web/20110803082104/http://www.nytimes.com/2006/10/25/dining/25fruit.html |archive-date=3 August 2011}}</ref>

=== Pests and diseases ===

[[File:Madige-Apfel-Frucht.jpg|thumb|Codling moth larva tunnelling inside an apple]]

A wide range of pests and diseases can affect the plant, including:

* 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.<ref>{{cite book |last=Jackson |first=H. S. |editor1-last=Lowther |editor1-first=Granville |editor2-last=Worthington |editor2-first=William |title=The Encyclopedia of Practical Horticulture: A Reference System of Commercial Horticulture, Covering the Practical and Scientific Phases of Horticulture, with Special Reference to Fruits and Vegetables |date=1914 |publisher=The Encyclopedia of Horticulture Corporation |pages=475–476 |chapter-url=https://archive.org/details/CUbiodiversity628875-8748/page/475 |access-date=1 August 2024 |volume=I |chapter=Powdery Mildew}}</ref> * 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).<ref>{{cite book |editor1-last=Lowther |editor1-first=Granville |editor2-last=Worthington |editor2-first=William |title=The Encyclopedia of Practical Horticulture: A Reference System of Commercial Horticulture, Covering the Practical and Scientific Phases of Horticulture, with Special Reference to Fruits and Vegetables |date=1914 |publisher=The Encyclopedia of Horticulture Corporation |location=North Yakima, Washington |pages=45–51 |url=https://archive.org/details/CUbiodiversity628875-8748/page/475 |access-date=1 August 2024 |volume=I }}</ref> 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.<ref>{{cite book |editor1-last=Coli |editor1-first=William M. |editor2-last=Los |editor2-first=Lorraine M. |year=2003 |chapter=Insect Pests |chapter-url=https://web.archive.org/web/20070221120428/http://www.umass.edu/fruitadvisor/NEAPMG/28-46.pdf |title=2003-2004 New England Apple Pest Management Guide |publisher=University of Massachusetts Amherst |pages=28–29 |url=http://www.umass.edu/fruitadvisor/NEAPMG/index.htm |access-date=3 March 2008 |archive-date=12 February 2008 |archive-url=https://web.archive.org/web/20080212130013/http://www.umass.edu/fruitadvisor/NEAPMG/index.htm |url-status=deviated }}</ref> * 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.<ref name="Bradley">{{cite book |last1=Atthowe |first1=Helen |last2=Gilkeson |first2=Linda A. |last3=Kite |first3=L. Patricia |last4=Michalak |first4=Patricia S. |last5=Pleasant |first5=Barbara |last6=Reich |first6=Lee |last7=Scheider |first7=Alfred F. |editor1-last=Bradley |editor1-first=Fern Marshall |editor2-last=Ellis |editor2-first=Bardara W. |editor3-last=Martin |editor3-first=Deborah L. |date=2009 |title=The Organic Gardener's Handbook of Natural Pest and Disease Control |url=https://archive.org/details/organicgardeners00brad_0/page/32 |url-access=limited |publisher=Rodale, Inc. |pages=32–34 |isbn=978-1-60529-677-7 |lccn=2009039996 |oclc=419860680}}</ref>

Among the most serious disease problems is a bacterial disease called fireblight, and three fungal diseases: ''Gymnosporangium'' rust, black spot,<ref>{{cite book |editor1-last=Coli |editor1-first=William M. |editor2-last=Berkett |editor2-first=Lorraine P. |editor3-last=Spitko |editor3-first=Robin |year=2003 |chapter=Other Apple Diseases |chapter-url=https://web.archive.org/web/20070221120311/http://www.umass.edu/fruitadvisor/NEAPMG/19-27.pdf |title=2003-2004 New England Apple Pest Management Guide |language=en |publisher=University of Massachusetts Amherst |pages=19–27 |url=http://www.umass.edu/fruitadvisor/NEAPMG/index.htm |access-date=3 March 2008 |archive-date=12 February 2008 |archive-url=https://web.archive.org/web/20080212130013/http://www.umass.edu/fruitadvisor/NEAPMG/index.htm |url-status=deviated }}</ref> and bitter rot.<ref>{{Cite journal |last1=Martin |first1=Phillip L. |last2=Krawczyk |first2=Teresa |last3=Khodadadi |first3=Fatemeh |last4=Aćimović |first4=Srđan G. |last5=Peter |first5=Kari A. |date=2021 |title=Bitter Rot of Apple in the Mid-Atlantic United States: Causal Species and Evaluation of the Impacts of Regional Weather Patterns and Cultivar Susceptibility |journal=Phytopathology |volume=111 |issue=6 |pages=966–981 |doi=10.1094/PHYTO-09-20-0432-R |issn=0031-949X |pmid=33487025 |s2cid=231701083 |doi-access=free|bibcode=2021PhPat.111..966M }}</ref> 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.<ref name="Bradley"/> 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.<ref>{{Cite journal |last=Erler |first=Fedai |date=1 January 2010 |title=Efficacy of tree trunk coating materials in the control of the apple clearwing, Synanthedon myopaeformis |journal=Journal of Insect Science |volume=10 |issue=1 |page=63 |doi=10.1673/031.010.6301 |pmc=3014806 |pmid=20672979}}</ref>

=== Cultivars === {{main|List of apple cultivars}} [[File:Golden Delicious, SweeTango, Granny Smith, and Gala apples 3.JPG|thumb|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.<ref>{{cite book |last1=Elzebroek |first1=A. T. G. |last2=Wind |first2=Koop |date=2008 |title=Guide to Cultivated Plants |url=https://books.google.com/books?id=YvU1XnUVxFQC&q=apple%20cultivars%207%2C500&pg=PT39 |url-status=live |language=en |location=Wallingford, United Kingdom |publisher=CABI |page=27 |isbn=978-1-84593-356-2 |lccn=2007028459 |oclc=156975183 |archive-url=https://web.archive.org/web/20201020231604/https://books.google.com/books?id=YvU1XnUVxFQC&q=apple%20cultivars%207,500&pg=PT39 |archive-date=20 October 2020 |access-date=6 October 2020}}</ref> Cultivars vary in their yield and the ultimate size of the tree, even when grown on the same rootstock.<ref name="England"/> Different cultivars are available for temperate and subtropical climates. The UK's National Fruit Collection in Kent includes over 2,000 apple cultivars.<ref>{{cite web |url=http://www.nationalfruitcollection.org.uk/index.php |title=Home|website=National Fruit Collection |access-date=2 December 2012 |url-status=live |archive-url=https://web.archive.org/web/20120615145659/http://www.nationalfruitcollection.org.uk/index.php |archive-date=15 June 2012 }}</ref> The University of Reading, responsible for developing the UK national collection database, provides access to search the national collection. Its work is part of the European Cooperative Programme for Plant Genetic Resources, with 38 countries participating in the Malus/Pyrus work group.<ref>{{cite web |url=http://www.ecpgr.cgiar.org/index.php?id=2501&tx_wfqbe_pi1[uid]=59 |title=ECPGR Malus/Pyrus Working Group Members |website=Ecpgr.cgiar.org |date=22 July 2002 |access-date=25 August 2014 |archive-url=https://web.archive.org/web/20140826114504/http://www.ecpgr.cgiar.org/index.php?id=2501&tx_wfqbe_pi1[uid& |archive-date=26 August 2014 }}</ref>

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.<ref name="apples1">{{cite web |url=http://casfs.ucsc.edu/publications/news%20and%20notes/Fall_06_N&N.pdf |archive-url=https://web.archive.org/web/20070811154017/http://casfs.ucsc.edu/publications/news%20and%20notes/Fall_06_N%26N.pdf |archive-date=11 August 2007 |title=Autumn Apple Musings |access-date=24 January 2008 |pages=1–2 |author=Tarjan, Sue |publisher=News & Notes of the UCSC Farm & Garden, Center for Agroecology & Sustainable Food Systems |date=Fall 2006 }}</ref>

In Europe, apple breeding programs are conducted at places such as Julius Kühn-Institut, the German federal research center for cultivated plants.<ref>{{cite journal |last1=Peil |first1=A. |last2=Dunemann |first2=F. |last3=Richter |first3=K. |last4=Hoefer |first4=M. |last5=Király |first5=I. |last6=Flachowsky |first6=H. |last7=Hanke |first7=M.-V. |date=2008 |title=Resistance Breeding in Apple at Dresden-Pillnitz |url=http://orgprints.org/13698/ |url-status=live |journal=Ecofruit - 13th International Conference on Cultivation Technique and Phytopathological Problems in Organic Fruit-Growing: Proceedings to the Conference from 18th February to 20th February 2008 at Weinsberg/Germany |language=de |pages=220–225 |archive-url=https://web.archive.org/web/20210128124933/https://orgprints.org/13698/ |archive-date=28 January 2021 |access-date=31 July 2024}}</ref>

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.<ref name=":0"/> Released by the University of Minnesota in 1991, the 'Honeycrisp' has become famous for its crispness and juiciness, thereby commanding high market prices.<ref name=":0">{{cite news |last1=Beck |first1=Kellen |title=How breeders bring out the best in new apples |url=https://mashable.com/article/apple-breeding |access-date=31 July 2024 |work=Mashable |date=17 October 2020 |archive-date=31 July 2024 |archive-url=https://web.archive.org/web/20240731195019/https://mashable.com/article/apple-breeding |url-status=live }}</ref> Unusually for a popular cultivar, the 'Honeycrisp' is not directly related to another popular apple cultivar but instead to two unsuccessful cultivars.<ref>{{cite news |last1=Migicovsky |first1=Zoë |title=How a few good apples spawned today's top varieties — and why breeders must branch out |url=https://theconversation.com/how-a-few-good-apples-spawned-todays-top-varieties-and-why-breeders-must-branch-out-164538 |access-date=31 July 2024 |work=The Conversation |date=22 August 2021 |archive-date=31 July 2024 |archive-url=https://web.archive.org/web/20240731195017/https://theconversation.com/how-a-few-good-apples-spawned-todays-top-varieties-and-why-breeders-must-branch-out-164538 |url-status=live }}</ref> However, it is also difficult to grow and to store, prompting the industry to seek hybrids that not only appeal to consumers but are also less costly for farmers to cultivate and last longer in storage.<ref name=":1">{{Cite news |last=Karp |first=David |date=November 3, 2015 |title=Beyond the Honeycrisp Apple |url=https://www.nytimes.com/2015/11/04/dining/beyond-the-honeycrisp-apple.html |archive-url=https://archive.today/20230815200005/https://www.nytimes.com/2015/11/04/dining/beyond-the-honeycrisp-apple.html |archive-date=August 15, 2023 |access-date=February 18, 2025 |work=The New York Times}}</ref> By the 2020s, about half of the new apple varieties entering the market in the United States and Canada are 'Honeycrisp' progeny.<ref>{{Cite news |last=Drost |first=Philip |date=November 16, 2024 |title=These aren't your Granny's Smiths: Why we have more apple varieties than ever before |url=https://www.cbc.ca/radio/costofliving/new-apple-varieties-1.7383868 |work=CBC News |department=Cost of Living}}</ref> 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.<ref name=":1"/>

thumb|Less common apple cultivars from an orchard in ItalyCommercially 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.<ref name="England">{{cite web |url=http://www.plantpress.com/wildlife/o523-apple.php |title=Apple – ''Malus domestica'' |access-date=22 January 2008 |publisher=Natural England |archive-url=https://web.archive.org/web/20080512000735/http://www.plantpress.com/wildlife/o523-apple.php |archive-date=12 May 2008}}</ref> Modern apples are generally sweeter than older cultivars, as popular tastes in apples have varied over time.<ref name=":2">{{Cite news |last=Ward |first=Rachel |date=November 20, 2018 |title=Goodbye, Red Delicious: Canada's favourite apples are now sweeter, more juicy |url=https://www.cbc.ca/news/canada/calgary/apple-best-kind-canada-1.4911742 |access-date=February 18, 2025 |work=CBC News}}</ref> Most North Americans and Europeans favor crunchy, sweet, and subacid apples.<ref name="Saba2020">{{cite news |last1=Saba |first1=Michel |date=September 11, 2020 |title=The McIntosh apple is a legend, but it could disappear from Quebec shops |url=https://montrealgazette.com/news/local-news/the-mcintosh-apple-is-a-legend-but-it-could-disappear-from-quebec-shops |archive-url=https://archive.today/20250219043550/https://www.montrealgazette.com/news/article506358.html |archive-date=February 19, 2025 |access-date=February 18, 2025 |work=The Montreal Gazette |agency=The Canadian Press}}</ref> Nevertheless, tart apples maintain a strong minority following.<ref name="World"/> In the United States today, the most popular apple varieties are the 'Ambrosia', 'Honeycrisp', and 'Jazz', according Nielsen data,<ref name=":1"/> while in Canada, the 'Honeycrisp', 'Ambrosia', and 'Gala' take the top spots.<ref name=":2"/> Together, these newer varieties have overtaken once dominant cultivars like the 'McIntosh' and the 'Red Delicious' in the North American market.<ref name=":2"/><ref name="Saba2020"/> Extremely sweet apples with barely any acid flavor are popular in Asia,<ref name="World">{{cite web |url=http://www.fas.usda.gov/htp2/circular/1998/98-03/applefea.html |title=World apple situation |access-date=24 January 2008 |archive-url=https://web.archive.org/web/20080211120700/http://www.fas.usda.gov/htp2/circular/1998/98-03/applefea.html |archive-date=11 February 2008 }}</ref> especially the Indian subcontinent.<ref name="apples1"/>

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.<ref name="Hobby Farms Mag.">{{cite magazine |last=Weaver |first=Sue |date=June–July 2003 |title=Crops & Gardening – Apples of Antiquity |magazine=Hobby Farms Magazine |url=http://www.hobbyfarms.com/apples-of-antiquity-3/ |url-status=live |archive-url=https://web.archive.org/web/20170219090902/http://www.hobbyfarms.com/apples-of-antiquity-3/ |archive-date=19 February 2017 }}</ref> A few old cultivars are still produced on a large scale, but many have been preserved by home gardeners and farmers who 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.<ref name="UofGeorgia">{{cite web |last1=Rieger |first1=Mark |title=Apple - ''Malus domestica'' |url=http://www.uga.edu/fruit/apple.html |website=HORT 3020: Intro Fruit Crops |access-date=22 January 2008 |publisher=University of Georgia |archive-url=https://web.archive.org/web/20080121045236/http://www.uga.edu/fruit/apple.html |archive-date=21 January 2008 }}</ref>

=== Production === {{Main|List of countries by apple production}} {{owidslider |start = 2024 |list = Template:OWID/Apple production#gallery |location = commons |caption = |title = |language = |file = link=|thumb|upright=1.6|Apple production |startingView = World }} {| class="wikitable floatright" style="width:12em; text-align:center;" |+ Apple production {{small|2023, millions of tonnes}} |- |{{CHN}}|| 49.6 |- |{{USA}}|| 5.2 |- |{{TUR}}|| 4.6 |- |{{POL}}|| 3.9 |- |{{IND}}|| 2.9 |- |{{IRN}}|| 2.2 |- |'''World''' || '''97.3''' |- |colspan=2 style="text-align: center;" |{{small|Source: FAOSTAT of the United Nations}}<ref name="faostat">{{cite web |date=2025 |title=Apple production in 2023; from pick lists: Crops/World Regions/Production Quantity |url=http://www.fao.org/faostat/en/#data/QC |access-date=18 February 2025 |publisher=FAOSTAT, UN Food and Agriculture Organization, Statistics Division}}</ref> |}

World production of apples in 2023 was 97 million tonnes, with China producing 51% of the total (table).<ref name=faostat/> Secondary producers were the United States, Turkey, and Poland.<ref name=faostat/>

== 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.<ref>{{cite book |last1=Nelson |first1=Lewis S. |last2=Shih |first2=Richard D. |last3=Balick |first3=Michael J. |author-link3=Michael Jeffrey Balick |date=2007 |title=Handbook of Poisonous and Injurious Plants |url=https://archive.org/details/handbookofpoison0000nels/page/211 |url-access=registration |language=en |edition=Second |location=New York |publisher=Springer for the New York Botanical Garden |pages=27, 211–212 |isbn=978-0387-31268-2 |lccn=2005938815 |oclc=77537459 |access-date=11 September 2024}}</ref> The U.S. National Library of Medicine's Hazardous Substances Data Bank records no cases of amygdalin poisoning from consuming apple seeds.<ref>{{cite web |title=Amygdalin |url=https://toxnet.nlm.nih.gov/cgi-bin/sis/search/a?dbs+hsdb:@term+@DOCNO+3559 |archive-url=https://web.archive.org/web/20170421094516/https://toxnet.nlm.nih.gov/cgi-bin/sis/search/a?dbs+hsdb:@term+@DOCNO+3559 |archive-date=21 April 2017 |access-date=20 April 2017 |publisher=Toxnet, US Library of Medicine}}</ref>

=== 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.<ref name="EU">{{cite web |title=General Information – Apple |url=http://foodallergens.ifr.ac.uk/food.lasso?selected_food=2#summary |archive-url=https://web.archive.org/web/20120723201454/http://foodallergens.ifr.ac.uk/food.lasso?selected_food=2 |archive-date=23 July 2012 |access-date=17 October 2011 |publisher=Informall}}</ref> 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,<ref name="EU" /> but in rare cases can also include life-threatening anaphylaxis.<ref>Landau, Elizabeth, [https://www.cnn.com/2009/HEALTH/04/08/oral.allergy.syndrome/index.html Oral allergy syndrome may explain mysterious reactions], 8 April 2009, ''CNN Health'', accessed 17 October 2011</ref> 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.<ref name="EU"/>

In other areas, such as the Mediterranean, some individuals have adverse reactions to apples because of their similarity to peaches.<ref name="EU"/> 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.<ref name="EU"/>

Breeding efforts have yet to produce a hypoallergenic fruit suitable for either of the two forms of apple allergy.<ref name="EU"/>

== Uses == {{See also|Cooking apple|Cider apple}}

=== Nutrition === {{Infobox nutritional value |name = Apples, with skin (edible parts) |water = 85.56 g |kJ = 218 |protein = 0.26 g |fat = 0.17 g |carbs = 13.81 g |fiber = 2.4 g |sugars = 10.39 |calcium_mg = 6 |iron_mg = 0.12 |copper_mg = 0.027 |magnesium_mg = 5 |phosphorus_mg = 11 |potassium_mg = 107 |sodium_mg = 1 |zinc_mg = 0.04 |manganese_mg = 0.035 |vitC_mg = 4.6 |thiamin_mg = 0.017 |riboflavin_mg = 0.026 |niacin_mg = 0.091 |pantothenic_mg = 0.061 |vitB6_mg = 0.041 |folate_ug = 3 |vitA_ug = 3 |betacarotene_ug = 27 |lutein_ug = 29 |vitE_mg = 0.18 |vitK_ug = 2.2 |note = [https://fdc.nal.usda.gov/food-details/1102644/nutrients Link to Full Nutrient Report of USDA FoodData Central] }}

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 {{cvt|100|g}} provides 52 calories and a moderate content of dietary fiber (table). Otherwise, there is low content of micronutrients, which are all below 10% of the Daily Value (table).

=== Culinary ===

thumb|left|Machine for paring, coring, and slicing apples, from Henry B. Scammell's 1897 handbook ''Cyclopedia of Valuable Receipts''

Apple varieties can be grouped as cooking apples, eating apples, and cider apples, the last so astringent as to be "almost inedible".<ref name="Davidson 2014"/> Apples are consumed as juice, raw in salads, baked in pies, cooked into sauces and apple butter, or baked.<ref>{{cite book |last1=Traverso |first1=Amy |others=Photographs by Squire Fox |date=2011 |title=The Apple Lover's Cookbook |url=https://archive.org/details/appleloverscookb0000trav |url-access=limited |language=en |edition=First |location=New York |publisher=W.W. Norton & Company |pages=16, 32, 35, 45, 92, 137, 262–263, 275 |isbn=978-0-393-06599-2 |lccn=2011016560 |oclc=711051767 |ol=16450839W}}</ref> They are sometimes used as an ingredient in savory foods, such as sausage and stuffing.<ref>{{Cite web |last=Kellogg |first=Kristi |date=15 January 2015 |title=81 Best Apple Recipes: Dinners, Desserts, Salads, and More |url=https://www.epicurious.com/ingredients/apple-of-my-eye-gallery |url-status=live |archive-url=https://web.archive.org/web/20201018215038/https://www.epicurious.com/ingredients/apple-of-my-eye-gallery |archive-date=18 October 2020 |access-date=17 October 2020 |website=Epicurious}}</ref>

Several techniques are used to preserve apples and apple products. Traditional methods include drying and making apple butter.<ref name="Davidson 2014"/> Juice and cider are produced commercially; cider is a significant industry in regions such as the West of England and Normandy.<ref name="Davidson 2014"/>

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.<ref>{{cite book |last1=Davidson |first1=Alan |author-link=Alan Davidson (food writer) |editor-last1=Jaine |editor-first1=Tom |editor-link1=Tom Jaine |others=Illustrated by Soun Vannithone |date=2014 |chapter=Toffee Apple |chapter-url=https://archive.org/details/oxfordcompaniont0000davi_w2w6/page/824/ |chapter-url-access=registration |title=The Oxford Companion to Food |language=en |edition=Third |location=Oxford |publisher=Oxford University Press |page=824 |isbn=978-0-19-967733-7 |lccn=2013957569 |oclc=890807357 |ol=27172691M |access-date=18 September 2024}}</ref><ref name="Lim 2012" /> Apples and honey are a ritual food pairing eaten during the Jewish New Year of Rosh Hashanah.<ref>{{cite web |last=Shurpin |first=Yehuda |title=Why All the Symbolic Rosh Hashanah Foods? "בולבול" |url=https://www.chabad.org/library/article_cdo/aid/3053586/jewish/Why-All-the-Symbolic-Rosh-Hashanah-Foods.htm |website=Chabad.org |access-date=21 March 2023 |archive-date=21 March 2023 |archive-url=https://web.archive.org/web/20230321154816/https://www.chabad.org/library/article_cdo/aid/3053586/jewish/Why-All-the-Symbolic-Rosh-Hashanah-Foods.htm |url-status=live }}</ref>

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.<ref name="Davidson 2014">{{cite book |last1=Davidson |first1=Alan |author-link=Alan Davidson (food writer) |editor-last1=Jaine |editor-first1=Tom |editor-link1=Tom Jaine |others=Illustrated by Soun Vannithone |date=2014 |chapter=Apple |chapter-url=https://archive.org/details/oxfordcompaniont0000davi_w2w6/page/27 |chapter-url-access=registration |title=The Oxford Companion to Food |language=en |edition=Third |location=Oxford |publisher=Oxford University Press |pages=27–31 |isbn=978-0-19-967733-7 |lccn=2013957569 |oclc=890807357 |ol=27172691M |access-date=18 September 2024}}</ref>

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.<ref name="Lim 2012" /> Through distillation, various alcoholic beverages can be produced, such as applejack, Calvados, and apple brandy.<ref name="Lim 2012" /><ref>{{cite book |last1=Yepsen |first1=Roger B. |date=2017 |orig-date=1994 |title=Apples |language=en |edition=Revised and Updated |location=New York |publisher=W.W. Norton & Company |page=52 |isbn=978-1-68268-019-3 |lccn=2017010136 |oclc=973918728}}</ref>

===Organic production=== Organic apples are commonly produced in the United States.<ref name=organic>{{cite web |url=http://www.agmrc.org/commodities-products/fruits/apples/organic-apples/ |title=Organic apples |access-date=23 February 2017 |publisher=USDA Agricultural Marketing Service |date=February 2016 |archive-url=https://web.archive.org/web/20170224052612/http://www.agmrc.org/commodities-products/fruits/apples/organic-apples/ |archive-date=24 February 2017 }}</ref> Due to infestations by key insects and diseases, organic production is difficult in Europe.<ref name="croplife">{{cite web |url=https://croplife.org/wp-content/uploads/pdf_files/European-Organic-apple-production-demonstrates-the-value-of-pesticides.pdf |title=European Organic Apple Production Demonstrates the Value of Pesticides |date=December 2011 |publisher=CropLife Foundation, Washington, DC |access-date=23 February 2017 |url-status=live |archive-url=https://web.archive.org/web/20170224054210/https://croplife.org/wp-content/uploads/pdf_files/European-Organic-apple-production-demonstrates-the-value-of-pesticides.pdf |archive-date=24 February 2017 }}</ref> 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.<ref name=croplife/> A light coating of kaolin, which forms a physical barrier to some pests, also may help prevent apple sun scalding.<ref name="Polomski & Reighard" />

=== Non-browning apples === Apple skins and seeds contain polyphenols.<ref name=Ribeiro2014>{{cite journal |last1=Ribeiro |first1=Flávia A.P. |last2=Gomes de Moura |first2=Carolina F. |last3=Aguiar |first3=Odair |last4=de Oliveira |first4=Flavia |last5=Spadari |first5=Regina C. |last6=Oliveira |first6=Nara R.C. |last7=Oshima |first7=Celina T.F. |last8=Ribeiro |first8=Daniel A. |title=The chemopreventive activity of apple against carcinogenesis: antioxidant activity and cell cycle control |journal=European Journal of Cancer Prevention |volume=23 |issue=5 |pages=477–480 |date=September 2014 |pmid=24366437 |doi=10.1097/CEJ.0000000000000005 |s2cid=23026644 |type=Review}}</ref> 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.<ref name="nic">{{Cite journal |last1=Nicolas |first1=J. J. |last2=Richard-Forget |first2=F. C. |last3=Goupy |first3=P. M. |last4=Amiot |first4=M. J. |last5=Aubert |first5=S. Y. |date=1 January 1994 |title=Enzymatic browning reactions in apple and apple products |journal=Critical Reviews in Food Science and Nutrition |volume=34 |issue=2 |pages=109–157 |doi=10.1080/10408399409527653 |pmid=8011143}}</ref> 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.<ref>{{cite web |title=PPO silencing |work=OK Specialty Fruits |publisher=Okanagan Specialty Fruits |date=2019 |url=https://www.okspecialtyfruits.com/our-science/ppo-silencing/ |access-date=14 November 2019 |url-status=live |archive-date=27 April 2021 |archive-url=https://web.archive.org/web/20210427083033/https://www.okspecialtyfruits.com/our-science/ppo-silencing/ }}</ref><ref>{{cite web |title=United States: GM non-browning Arctic apple expands into foodservice |url=https://www.freshfruitportal.com/news/2019/08/13/u-s-gm-non-browning-arctic-apple-expands-into-foodservice/ |publisher=Fresh Fruit Portal |access-date=14 November 2019 |date=13 August 2019 |archive-url=https://web.archive.org/web/20210627220702/https://www.freshfruitportal.com/news/2019/08/13/u-s-gm-non-browning-arctic-apple-expands-into-foodservice/ |url-status=live |archive-date=27 June 2021}}</ref> 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.<ref name="fda2015">{{cite web |url=https://www.fda.gov/Food/IngredientsPackagingLabeling/GEPlants/Submissions/ucm436163.htm |title=Okanagan Specialty Fruits: Biotechnology Consultation Agency Response Letter BNF 000132 |website=U.S. Food and Drug Administration |date=March 20, 2015 |access-date=14 November 2019 |archive-date=31 October 2017 |archive-url=https://wayback.archive-it.org/7993/20171031091829/https://www.fda.gov/Food/IngredientsPackagingLabeling/GEPlants/Submissions/ucm436163.htm}}</ref><ref name="cfia">{{cite web |title=Questions and answers: Arctic Apple |publisher=Canadian Food Inspection Agency, Government of Canada |url=http://www.inspection.gc.ca/plants/plants-with-novel-traits/general-public/questions-and-answers-arctic-apple/eng/1426884802194/1426884861294 |access-date=14 November 2019|date=8 September 2017 |url-status=live |archive-date=19 September 2018 |archive-url=https://web.archive.org/web/20180919152434/http://www.inspection.gc.ca/plants/plants-with-novel-traits/general-public/questions-and-answers-arctic-apple/eng/1426884802194/1426884861294}}</ref>

=== Other products === Apple seed oil is obtained by pressing apple seeds for manufacturing cosmetics.<ref>{{cite journal |doi=10.2202/1556-3758.1283 |title=Proximate Composition of the Apple Seed and Characterization of Its Oil |year=2007 |last1=Yu |first1=Xiuzhu |last2=Van De Voort |first2=Frederick R. |last3=Li |first3=Zhixi |last4=Yue |first4=Tianli |journal=International Journal of Food Engineering |volume=3 |issue=5|s2cid=98590230 }}</ref>

== In culture == {{Main|Apple (symbolism)}}

=== Germanic paganism === [[File:Carl Larsson Brita as Iduna.jpg|upright|thumb|alt=Illustration of girl in a red dress, holding 3 candles in one hand and a basket of apples in the other|"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.<ref name="Davidson 1990">{{cite book |last1=Davidson |first1=Hilda Roderick Ellis |author-link=Hilda Ellis Davidson |date=1990 |orig-date=1st pub. 1964 |title=Gods and Myths of Northern Europe |language=en |location=London |publisher=Penguin Books |pages=165–166 |isbn=0-14-013627-4 |oclc=29336401}}</ref>

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.<ref name="Davidson 1990" /> 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.<ref>{{cite book |last1=Davidson |first1=Hilda Ellis |author-link=Hilda Ellis Davidson |date=1998 |title=Roles of the Northern Goddess |language=en |location=London; New York |publisher=Routledge |pages=146–147 |doi=10.4324/9780203025550 |isbn=0-415-13610-5 |lccn=97018309 |oclc=48138055}}</ref>

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."<ref name="Davidson 1990" />

=== Greek mythology ===

[[File:Hercules Musei Capitolini MC1265 n2.jpg|thumb|upright=0.6|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.<ref>{{cite book |last1=Biedermann |first1=Hans |translator-last1=Hulbert |translator-first1=James |date=1992 |title=Dictionary of Symbolism |url=https://archive.org/details/dictionaryofsymb00bied_0/page/16 |url-access=registration |language=en |location=New York |publisher=Facts on File |pages=16–17 |isbn=978-0-8160-2593-0 |lccn=91044933 |oclc=25092926 |access-date=3 October 2024}}</ref> 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.<ref name="Ruck 2001">{{cite book |last1=Ruck |first1=Carl A. P. |author-link1=Carl A. P. Ruck |last2=Staples |first2=Blaise D. |author-link2=Blaise Daniel Staples |last3=Heinrich |first3=Clark |author-link3=Clark Heinrich |date=2001 |title=The apples of Apollo: pagan and Christian mysteries of the Eucharist |language=en |edition= |location=Durham, North Carolina |publisher=Carolina Academic Press |pages=64–70 |isbn=978-0-89089-924-3 |lccn=00040351 |oclc=46337324}}</ref>

The Greek goddess of discord, Eris, became disgruntled after she was excluded from the wedding of Peleus and Thetis.<ref>{{cite web |title=Eris - Greek Goddess of Strife & Discord (Roman Discordia) |url=https://www.theoi.com/Daimon/Eris.html |website=Theoi Project |publisher=Aaron J. Atsma |access-date=26 September 2024 |archive-url=https://web.archive.org/web/20240925003522/https://www.theoi.com/Daimon/Eris.html |archive-date=25 September 2024 |language=en |url-status=live}}</ref> 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.<ref>{{cite book |last1=Lucian |author-link1=Lucian |translator-last1=Fowler |translator-first1=H.W. |translator-link1=H. W. Fowler |translator-last2=Fowler |translator-first2=F.G. |translator-link2= Francis George Fowler |date=1905 |title=The Works of Lucian of Samosata |url=https://archive.org/details/worksoflucianofs01luciuoft/page/78 |language=en |volume=I |edition=First |location=Oxford |publisher=Clarendon Press |pages=78–85 |lccn=06001045 |oclc=506365 |access-date=26 September 2024}}</ref><ref>{{cite web |title=Judgement of Paris - Greek Mythology |url=https://www.theoi.com/Olympios/JudgementParis.html |website=Theoi Project |publisher=Aaron J. Atsma |access-date=26 September 2024 |archive-url=https://web.archive.org/web/20240824120807/https://www.theoi.com/Olympios/JudgementParis.html |archive-date=24 August 2024 |language=en |url-status=live}}</ref>

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:<ref>{{cite book |last1=Plato |author-link1=Plato |editor-last1=Cooper |editor-first1=John M. |editor-last2=Hutchinson |editor-first2=D.S. |translator-last1=Edmonds |translator-first1=J.M. |translator-last2=Cooper |translator-first2=John M. |date=1997 |chapter=Epigrams |title=Complete Works |chapter-url=https://archive.org/details/completeworks00plat/page/1744 |chapter-url-access=registration |language=en |location=Indianapolis, Indiana |publisher=Hackett Publishing |page=1744 |isbn=0-87220-349-2 |lccn=96053280 |oclc=36178550 |access-date=27 September 2024}}</ref>

{{blockquote | text = 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. | sign = Plato | source = 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),<ref name="Ruck 2001"/> 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.<ref>{{cite book |last1=Pinsent |first1=John |author-link1=John Pinsent |date=1969 |title=Greek Mythology |url=https://archive.org/details/greekmythology00pins/page/79 |url-access=registration |language=en |edition=First |location=London |publisher=Paul Hamlyn |page=79 |isbn=978-0-600-02422-4 |lccn=78449216 |oclc=61702 |access-date=3 October 2024}}</ref><ref>{{cite web |title=Atalanta (Atalante) - Arcadian Heroine of Greek Mythology |url=https://www.theoi.com/Heroine/Atalanta.html |website=Theoi Project |publisher=Aaron J. Atsma |access-date=3 October 2024 |archive-url=https://web.archive.org/web/20240927101406/https://www.theoi.com/Heroine/Atalanta.html |archive-date=27 September 2024 |language=en |url-status=live}}</ref>

=== 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".<ref>{{cite book |last1=Flieger |first1=Verlyn |author-link1=Verlyn Flieger |date=2005 |title=Interrupted Music: The Making of Tolkien's Mythology |language=en |location=Kent, Ohio |publisher=Kent State University Press |pages=122–123 |isbn=978-0-87338-824-5 |lccn=2004024490 |oclc=56805947}}</ref>

=== China === thumb|''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.<ref name="Lisa Lim" /><ref name="English in China" /> 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.<ref name="English in China">{{cite web |date=22 December 2019 |title=Why Do the Chinese Give Apples Around Christmas? |url=https://teach-english-in-china.co.uk/why-do-the-chinese-give-apples-around-christmas/ |url-status=live |archive-url=https://web.archive.org/web/20201001003304/https://teach-english-in-china.co.uk/why-do-the-chinese-give-apples-around-christmas/ |archive-date=1 October 2020 |access-date=3 September 2024 |website=Teach English In China}}</ref>

=== Christian art === [[File:Albrecht Dürer - Adam and Eve (Prado) 2.jpg|thumb|''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.<ref name="Macrone">{{cite book |last1=Macrone |first1=Michael |title=Brush up your Bible! |date=1998 |publisher=Gramercy Books |location=New York |isbn=978-0-517-20189-3 |oclc=38270894 |pages=15–16, 340–341 |url=https://archive.org/details/brushupyourbible00macr |access-date=31 July 2024}}</ref> 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''.<ref>{{cite book |last1=Kissling |first1=Paul J. |date=2004 |title=Genesis |url=https://books.google.com/books?id=lotBnvqdmeQC&q=Genesis+apple&pg=PA193 |url-status=live |language=en |volume=1 |location=Joplin, Missouri |publisher=College Press |page=193 |isbn=978-0-89900-875-2 |lccn=2004022577 |oclc=56672257 |archive-url=https://web.archive.org/web/20210126053142/https://books.google.com/books?id=lotBnvqdmeQC&q=Genesis+apple&pg=PA193 |archive-date=26 January 2021 |access-date=6 October 2020}}</ref> The tree of the forbidden fruit is called "the tree of the knowledge of good and evil" in Genesis 2:17,<ref>{{bibleverse||Genesis|2:17|ESV}}</ref> and the Latin for "good and evil" is ''bonum et malum''.<ref>{{cite book |last1=Hendel |first1=Ronald S. |date=2013 |title=The Book of Genesis: A Biography |url=https://books.google.com/books?id=xBPpIHwcZMUC&q=Genesis+apple+malum&pg=PA114 |url-status=live |language=en |location=Princeton, New Jersey |publisher=Princeton University Press |page=114 |isbn=978-0-69114012-4 |lccn=2012015634 |oclc=788265521 |archive-url=https://web.archive.org/web/20230305222115/https://books.google.com/books?id=xBPpIHwcZMUC&q=Genesis+apple+malum&pg=PA114#v=snippet&q=Genesis%20apple%20malum&f=false |archive-date=5 March 2023 |access-date=4 October 2024}} </ref>

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.<ref name="Macrone"/>

=== 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".<ref>{{cite book |editor-last1=Mieder |editor-first1=Wolfgang |editor-link1=Wolfgang Mieder |editor-last2=Kingsbury |editor-first2=Stewart A. |editor-last3=Harder |editor-first3=Kelsie B. |editor-link3=Kelsie B. Harder |date=1996 |orig-date=1992 |title=A Dictionary of American Proverbs |url=https://archive.org/details/dictionaryofamer00wolf_0/page/23 |url-access=registration |language=en |edition=Paperback |location=New York |publisher=Oxford University Press |page=23 |isbn=978-0-19-511133-0 |lccn=91015508 |oclc=23693799 |access-date=23 August 2024}}</ref> 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.<ref>{{cite book |last1=Pollan |first1=Michael |author1-link=Michael Pollan |date=2001 |title=The Botany of Desire: A Plant's-Eye View of the World |url=https://archive.org/details/botanyofdesirepl0000poll_v5w7 |url-access=limited |language=en |edition=First |location=New York |publisher=Random House |pages=9, 22, 50 |isbn=978-0-375-50129-6 |lccn=00066479 |oclc=49803415}}</ref>

== See also ==

* Apple chip * Apple cider * Apple juice * Applecrab, apple–crabapple hybrids for eating * Isaac Newton's apple tree * Johnny Appleseed * List of apple dishes

== References == {{reflist}}

== Further reading == * {{cite book |last1=Browning |first1=Frank |author1-link=Frank Browning (author) |date=1998 |title=Apples |url=https://archive.org/details/apples00brow |url-access=registration |edition=First |location=New York |publisher=North Point Press |isbn=978-0-86547-537-3 |lccn=98027252 |oclc=39235786 |ref=none}} * {{cite book |editor-last1=Hanson |editor-first1=Beth |editor-last2=Marinelli |editor-first2=Janet |editor-last3=Saphire |editor-first3=Sigrun Wolff |editor-last4=Tebbitt |editor-first4=Mark |date=2003 |title=The Best Apples to Buy and Grow |url=https://archive.org/details/bestapplestobuyg0000unse |url-access=registration |language=en |edition=First |location=Brooklyn, New York |publisher=Brooklyn Botanic Garden |isbn=978-1-889538-66-2 |oclc=60384060 |ref=none}} * {{cite book |last1=Juniper |first1=Barrie E. |last2=Mabberley |first2=David J. |author1-link=Barrie Juniper |author2-link=David Mabberley |date=2006 |title=The Story of the Apple |url=https://archive.org/details/storyofapple0000juni |url-access=registration |edition=First |location=Portland, Oregon |publisher=Timber Press |isbn=978-0-88192-784-9 |lccn=2006011869 |oclc=67383484 |ref=none}} * {{cite book |last1=Phillips |first1=Michael |date=1998 |title=The Apple Grower: A Guide for the Organic Orchardist |url=https://archive.org/details/applegrowerguide0000phil |url-access=registration |language=en |edition=First |location=White River Junction, Vermont |publisher=Chelsea Green Publishing |isbn=978-1-890132-04-0 |lccn=98003631 |oclc=38731995 |ref=none}} * {{cite book |last1=Sanders |first1=Rosie |title=The Apple Book |date=2010 |publisher=Frances Lincoln Limited |location=London |isbn=978-0-7112-3141-2 |edition=Second |language=en |oclc=646397065 |ref=none}}

== External links ==

* {{Commons category-inline|Apples}} * {{Cookbook-inline|Apple}}

* Category:Fruits originating in Asia * Category:Plants described in 1768