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{{Short description|Fruit that grows on a tree}}
{{otheruses4|the fruit|the computer corporation|Apple Inc|other uses|Apple (disambiguation)}}
{{About|the fruit|the technology company|Apple Inc.|other uses}}
{{Taxobox
{{Redirect|Apple tree}}
| color = lightgreen
{{pp-semi-indef}}
| name = Apple
{{pp-move}}
| image = Koeh-108.jpg
{{Use dmy dates|date=September 2019}}
| image_width = 240px
{{Good article}}
| image_caption = Apple tree (''Malus domestica'')
{{Speciesbox
| regnum = ]ae
|name = Apple
| divisio = ]
|image = Pink lady and cross section.jpg
| classis = ]
| ordo = ] |image_caption = ']' apples
|image2 = Malus domestica a1.jpg{{!}}class=notpageimage <!-- See ] -->
| familia = ]
|image2_caption = Flowers
| subfamilia = ]
| genus = '']'' |genus = Malus
| species = '''''M. domestica''''' |species = domestica
|authority = (]) ]
| binomial = ''Malus domestica''
|synonyms =
| binomial_authority = ]
* ''M.&nbsp;communis'' <small>Desf., 1768</small>
* ''M.&nbsp;pumila'' <small>Mil.</small>
* ''M.&nbsp;frutescens'' <small>Medik.</small>
* ''M.&nbsp;paradisiaca'' <small>(L.) Medikus</small>
* ''M.&nbsp;sylvestris'' <small>Mil.</small>
* ''Pyrus malus'' <small>L.</small>
* ''Pyrus malus'' var. ''paradisiaca'' <small>L.</small>
* ''Pyrus dioica'' <small>Moench</small>
|synonyms_ref =<ref name="FNA" /><ref name="POWO" />
}} }}
]'' apple in ]]]
]


An '''apple''' is a round, edible ] produced by an '''apple tree''' (], among them the '''domestic''' or '''orchard apple'''; '''''Malus domestica'''''). Apple ] are ] worldwide and are the most widely grown species in the ] '']''. The ] originated in ], where its wild ancestor, '']'', is still found. Apples have been grown for thousands of years in ] and were introduced to ] by ]. Apples have ] and ] significance in many cultures, including ], ], and ] tradition.
The '''apple''' is the ] ] of the apple tree, ] ''Malus domestica'' in the ] family ]. It is one of the most widely ] tree fruits. The tree is small and ], reaching 5-12 m tall, with a broad, often densely twiggy crown.


Apples grown from seed tend to be very different from those of their parents, and the resultant fruit frequently lacks desired characteristics. For commercial purposes, including botanical evaluation, apple ]s are propagated by clonal ] onto ]s. 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.
The ] are ] simple ovals 5-12 cm long and 3-6 cm broad on a 2-5 cm ] with an acute tip, serrated margin and a slightly downy underside. ]s are produced in ]simultaneous with the budding of the leaves.


There are ]. Different cultivars are bred for various tastes and uses, including ], eating raw, and ] or ] production. Trees and fruit are prone to ], bacterial, and pest problems, which can be controlled by a number of ] and non-organic means. In 2010, the fruit's ] was ] as part of research on disease control and selective breeding in apple production.
The flowers are white with a pink tinge that gradually fades, five ]ed, 2.5-3.5 ] in diameter. The fruit matures in ], and is typically 5-9 cm diameter. The centre of the fruit contains five ]s arranged in a ], each carpel containing one to three ]s.


== Etymology ==
==Botanical information==
The word ''apple'', whose ] ancestor is {{lang|ang-Latn|æppel}}, is descended from the ] noun {{lang|gem-x-proto|*aplaz}}, descended in turn from ] {{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=] |language=en |location=Hong Kong, China |publisher=] |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 ]s. This can be compared to the 14th-century ] expression {{lang|enm|appel of paradis}}, meaning a ].<ref>{{cite web |title=Origin and meaning of "apple" by Online Etymology Dictionary |url=https://www.etymonline.com/word/apple |website=] |access-date=22 November 2019 |archive-date=21 December 2019 |archive-url=https://web.archive.org/web/20191221020212/https://www.etymonline.com/word/apple |url-status=live }}</ref>
The wild ] of ''Malus domestica'' is '']''. It has no common name in English, but is known in ], where it is native, as 'alma'; in fact, the region where it is thought to originate is called ], or 'father of the apples'. This tree is still found wild in the mountains of ] in southern Kazakhstan, ], ], and ], ].


== Description ==
For many years, there was a debate about whether ''M. domestica'' evolved from chance hybridisation among various wild species. Recent DNA analysis by Barrie Juniper, Emeritus Fellow in the Department of Plant Sciences at ] and others, has indicated, however, that the hybridisation theory is probably false. Instead, it appears that a single species still growing in the Ili Valley on the northern slopes of the Tien Shan mountains at the border of northwest China and the former Soviet Republic of Kazakhstan is the progenitor of the apples we eat today. Leaves taken from trees in this area were analyzed for DNA composition, which showed them all to belong to the species '']'', with some genetic sequences common to ''M. domestica''.{{Fact|date=February 2007}}


The apple is a ] tree, generally standing {{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=] |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 ] 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=] |page=909 |isbn=978-1-930723-84-9 |oclc=859541992 |access-date=27 July 2024}}</ref>
Some individual ''M. sieversii'', recently planted by the US government at a research facility, resist many ] that affect domestic apples, and are the subject of continuing research to develop new disease-resistant apples.


The buds are egg-shaped and dark red or purple in color; they range in size from 3 to 5&nbsp;millimeters, but are usually less than 4&nbsp;mm. The ]s 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 (]), medium or wide in width, somewhat egg-shaped with the wider portion toward their base (]), or even with sides that are more parallel to each other instead of curved (]) 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 ] by short leaf stems {{cvt|1 to 3.5|cm||sp=us|adj=on|frac=2}} long.<ref name="NC Extension" /><ref name="FNA" />
Other ] that were previously thought to have made contributions to the ] of the domestic apples are '']'' and '']'', but there is no hard evidence for this in older apple ]s. These and other ''Malus'' species have been used in some recent breeding programmes to develop apples suitable for growing in climates unsuitable for ''M. domestica'', mainly for increased cold tolerance.{{Fact|date=February 2007}}


] are produced in ] simultaneously with the budding of the leaves and are produced on spurs and some long ]s.<ref name="UofGeorgia" /> When the flower buds first begin to open the ]s 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-<!--hyphen is vital here: not five flowers, but one 5-petaled flower-->petaled flowers are group in an ] consisting of a ] 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=] |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=] |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" />
The apple tree was perhaps the earliest tree to be cultivated, and apples have remained an important food in all cooler climates. To a greater degree than other tree fruit, except possibly ], apples store for months while still retaining much of their nutritive value. Winter apples, picked in late autumn and stored just above freezing, have been an important food in ] and ] for millennia, as well as in ] and in the ] since the ].{{Fact|date=February 2007}}


{{gallery|mode=packed
==Apple cultivars==
|Apple Blossom @ Manali.jpg|Apple blossoms
{{refimprove|date=July 2007}}
|Koeh-108.jpg|Botanical illustration
]
}}
] ]]
:''See ] for a listing.''
There are more than 7,500 known ]s of apples. Different cultivars are available for ] and ] climates. Reputedly the world's biggest collection of apple ]s is housed at the in ]. Apples do not flower in tropical climates because they have a chilling requirement.


=== Fruit ===
Commercially popular apple cultivars are soft but crisp. Other desired qualities in modern commercial apple breeding are a colourful skin, absence of ]ing, ease of shipping, lengthy storage ability, high yields, disease resistance, typical "]" apple shape, long stem (to allow ]s to penetrate the top of the fruit), and popular flavour. The taste of the apple doesn't depend on the colour. If the apple is bright red it could still taste bad.


The ] is a ] that matures in late ] or ].<ref name="FNA" /> The true fruits or ]s 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=] |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>
Old cultivars are often oddly shaped, russeted, and have a variety of textures and colours. Many of them have excellent flavour (often better than most modern cultivars), but may have other problems which make them commercially unviable, such as low yield, liability to disease, or poor tolerance for storage or transport. A few old cultivars are still produced on a large scale, but many have been kept alive by home gardeners and farmers that sell directly to local markets. Many unusual and locally important cultivars with their own unique taste and appearance are out there to discover; apple conservation campaigns have sprung up around the world to preserve such local cultivars from extinction. In the ] old cultivars such as Cox's Orange Pippin and Egremont Russett are still commercially important even though by modern standards they are low yielding and disease prone.


<gallery class=center mode=nolines widths=400 heights=250>
Although most cultivars are bred for eating fresh (dessert apples), some are cultivated specifically for cooking (]s) or producing ]. ]s are typically too tart and astringent to eat fresh, but they give the beverage a rich flavour that dessert apples cannot.
File:Apple anatomy, flower and fruit compared.svg|How apple fruit derives from flower structures
</gallery>


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=] |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"/>
Modern apples are, as a rule, sweeter than older cultivars. Most ]ns and ]ans favour sweet, subacid apples, but tart apples have a strong minority following. Extremely sweet apples with barely any acid flavour are popular in ] and especially India.


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=] |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>
Tastes in apples vary from one person to another and change continually over time. As an example, the ] of ] made its reputation for apple growing on Red Delicious. In recent years, many apple connoisseurs have come to regard the Red Delicious as inferior to cultivars such as ] and ] due to its merely mild flavour and insufficiently firm texture.


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 |language=en |volume=I: Tree and Tropical Fruits |location=New York |publisher=] |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 ], making it rough and brown. The skin is covered in a protective layer of ].<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 |url-status=dead |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" />
==Growing apples==
===Apple breeding===
]
]
]


<gallery mode=packed caption="Apples can have any amount of overcolor, a darker tint over a pale groundcolor.">
Like most perennial fruits, apples ordinarily propagate asexually by ]. Seedling apples are different from their parents, sometimes radically. Most new apple cultivars originate as seedlings, which either arise by chance or are bred by deliberately crossing cultivars with promising characteristics. The words 'seedling', 'pippin', and 'kernel' in the name of an apple cultivar suggest that it originated as a seedling. Apples can also form ]s (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.
File:Yellow Transparent (cropped).jpg|0% overcolor
File:Lobo (apple) (cropped).jpg|100% overcolor
</gallery>


=== Chemistry ===
Some breeders have crossed ordinary apples with ] or unusually hardy apples in order to produce ] cultivars. For example, the Excelsior Experiment Station of the ] has, since the 1930s, introduced a steady progression of important hardy apples that are widely grown, both commercially and by backyard orchardists, throughout ] and ]. Its most important introductions have included ']' (which is the most widely cultivated apple in Minnesota), 'Wealthy', 'Honeygold', and ']'. <ref> </ref>


Important volatile compounds in apples that contribute to their scent and flavour include ], ], ], ], 2-methylbutanal, ], ], ethyl 2-methylpropionate, ], ethyl 2-methyl butyrate, ], ], ], 2-methylbutyl acetate, 1-propyl butyrate, ], ], ], trans-2-hexenal, ], ].<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=] |date=January 1967 |volume=15 |issue=1 |page=29 |doi=10.1021/jf60149a032}}</ref>
===Pollination===
Apples are self-incompatible; they must ] to develop fruit. During the flowering each season, apple growers usually provide ]s to carry the pollen. ] hives are most commonly used. ]s are also used as supplemental pollinators in commercial orchards. ] ]s are sometimes present in orchards, but not usually in enough quantity to be significant pollinators.


== Taxonomy ==
There are four to seven pollination groups in apples depending on climate:
The apple as a species has more than 100 alternative scientific names, or ].<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.&nbsp;pumila'', ''M.&nbsp;communis'', ''M.&nbsp;frutescens'', and ''Pyrus dioica'' ( Rosaceae ) |journal=] |date=April 2010 |volume=59 |issue=2 |pages=650–652 |doi=10.1002/tax.592038 |language=en}}</ref> In modern times, ''Malus pumila'' and ''Malus domestica'' are the two main names in use. ''M.&nbsp;pumila'' is the older name, but ''M.&nbsp;domestica'' has become much more commonly used starting in the 21st century, especially in the western world. Two proposals were made to make ''M.&nbsp;domestica'' a ]: the earlier proposal was voted down by the Committee for Vascular Plants of the ] 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=] |volume=66 |issue=2 |pages=500–513 |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.&nbsp;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 }}</ref> Nevertheless, some works published after 2017 still use ''M.&nbsp;pumila'' as the ], 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 ] in 1753, the pears, apples, and quinces were combined into one genus that he named '']'' and he named the apple as ''Pyrus malus''. This was widely accepted, however the botanist ] published an alternate classification in ] with the apple species separated from ''Pyrus'' in 1754. He did not clearly indicate that by ''Malus pumila'' he meant the domesticated apple. Nonetheless, it was used as such by many botanists. When ] published his scientific description of the apple in 1803 it may have been a new combination of ''P.&nbsp;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 ] in 1786.<ref name="POWO" />
* Group A – Early flowering, May 1 to 3 in England (], ])
* Group B – May 4 to 7 (], ])
* Group C – Mid-season flowering, May 8 to 11 (], ])
* Group D – Mid/Late season flowering, May 12 to 15 (], ]).
* Group E – Late flowering, May 16 to 18 (], ])
* Group F – May 19 to 23 (])
* Group H – May 24 to 28 (])


=== Genome ===
One cultivar can be pollinized by a compatible cultivar from the same group or close (A with A or A with B but not A with C or D).
{{further |Apple genome}}
]
===Maturation and harvest===


Apples are ], with two sets of ]s per cell (though triploid cultivars, with three sets, are not uncommon), have 17 chromosomes and an estimated ] 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 ']'.<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=] |volume=3 |issue=1 |page=16057 |pmid=27917289 |pmc=5120355 |bibcode=2016HorR....316057D }}</ref> in part owing to the high degree of ] 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>
Cultivars vary in their yield and the ultimate size of the tree, even when grown on the same rootstock. Some cultivars, if left unpruned, will grow very large, which allows them to bear a great deal more fruit, but makes harvest very difficult. Mature trees typically bear 40-200 kg of apples each year, though productivity can be close to zero in poor years. Apples are harvested using three-point ladders that are designed to fit amongst the branches. Dwarf trees will bear about 10-80 kg of fruit per year.


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 '']'' following domestication.<ref name="Duan 2017"/>
===Pests and diseases===
{{Main|List of apple diseases}}
]
]


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


=== History ===
Among the most serious disease problems are ], a bacterial disease; and '']'' rust, ], and ], three fungal diseases.
]


]'' apple in Kazakhstan]]
The ] is the most serious insect pest. Others include ] and ].


] 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 '']'', found growing wild in the ] in southern ], ], ], and ].<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 |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 }}</ref> Cultivation of the species, most likely beginning on the forested flanks of the ] mountains, progressed over a long period of time and permitted secondary ] of genes from other species into the open-pollinated seeds. Significant exchange with '']'', the crabapple, resulted in populations of apples being more related to crabapples than to the more ] 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 |pages=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>
Young apple trees are also prone to mammal pests like mice and deer, which feed on the soft bark of the trees, especially in winter.


The apple is thought to have been domesticated 4,000–10,000 years ago in the ] mountains, and then to have travelled along the ] to Europe, with hybridization and introgression of wild crabapples from Siberia (''M.&nbsp;baccata''), the Caucasus (''M.&nbsp;orientalis''), and Europe (''M.&nbsp;sylvestris''). Only the ''M.&nbsp;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>
Organic apples are commonly produced in the United States.<ref>http://www.bestapples.com/facts/organic.html The Best Organic Apples On Earth</ref> Organic production is difficult in Europe, though a few orchards have done so with commercial success, using disease-resistant cultivars and the very best cultural controls. The latest tool in the organic repertoire is a spray of a light coating of ] clay, which forms a physical barrier to some pests, and also helps prevent apple sun scald.


Chinese soft apples, such as '']'' and '']'', have been cultivated as dessert apples for more than 2,000 years in China. These are thought to be hybrids between ''M.&nbsp;baccata'' and ''M.&nbsp;sieversii'' in Kazakhstan.<ref name="Duan 2017"/>
{{see also|List of Lepidoptera which feed on Malus}}


Among the traits selected for by human growers are size, fruit acidity, color, firmness, and soluble sugar. Unusually for domesticated fruits, the wild ''M.&nbsp;sieversii'' origin is only slightly smaller than the modern domesticated apple.<ref name="Duan 2017"/>
==Commerce==
]
]
At least 55 million tonnes of apples were grown worldwide in 2005, with a value of about $10 billion. ] produced about two-fifth of this total. ] is the second leading producer, with more than 7.5% of the world production. ], ], ] and ] are among the leading apple exporters.


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=]; 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>
In the ], more than 60% of all the apples sold commercially are grown in ] state. Imported apples from ] and other more temperate areas are competing with US production and increasing each year.


There is indirect evidence of apple cultivation in the third millennium BCE in the ].<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.
==Apples as food==
<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=] |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
Different cultivars of apples have a distinct different taste, and this can be separated into two separate factors of flavour and texture.
| filename = Wild Apples by Henry David Thoreau - read by Kevin S for LibriVox's Short Nonfiction Collection Vol. 065 (2019).ogg
| title = "Wild Apples"<br />by Henry David Thoreau<br /><small>Read by Kevin S for LibriVox</small>
| description = Audio 01:01:35 ()
| pos = right
| type = speech
| image = ]
}}


The Roman writer ] 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=] |location=London |page=303 |url=https://archive.org/details/naturalhistoryof03plin/page/303 |access-date=3 August 2024 |volume=III}}</ref> Though methods like this will extend the availabity of reasonably fresh apples, without refrigeration their lifespan is limited. Even sturdy winter apple varieties will only keep well until December in cool climates.<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=] |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 |last1=Adamson |first1=Melitta Weiss |date=2004 |title=Food in Medieval Times |url=https://archive.org/details/foodinmedievalti0000adam |url-access=limited |language=en |edition=First |location=Westport, Connecticut |publisher=] |pages=19–20 |isbn=978-0-313-32147-4 |lccn=2004014054 |oclc=55738647}}</ref>
Apples can be canned, juiced, and optionally fermented to produce ], ], ], ], and ]. Distilled apple cider produces the ] ] and ]. ] can also be made. They make a popular lunchbox fruit as well.


Of the many Old World plants that the Spanish introduced to ] 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=] |volume=77 |issue=4 |pages=661–677 |doi=10.4067/s0716-078x2004000400009 |language=es |doi-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 ] by Reverend ] 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=] |page=40 |lccn=62009906 |oclc=710612 |access-date=10 August 2024}}</ref> The only apples native to North America are ].<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=] |page=122 |isbn=978-0-920656-11-2 |oclc=1336124440 |access-date=10 August 2024}}</ref>
Apples are an important ingredient in many winter ]s, for example ], apple ], ] and ]. They are often eaten ] or ], and they can also be dried and eaten or re-constituted (soaked in water, alcohol or some other liquid) for later use. Puréed apples are generally known as ]. Apples are also made into ] and apple jelly. They are also used cooked in meat dishes.
*In the UK, a ] is a traditional confection made by coating an apple in hot ] and allowing it to cool. Similar treats in the US are ]s (coated in a hard shell of crystallised sugar syrup), and ], coated with cooled ].
*Apples are eaten with honey at the Jewish New Year of ] to symbolize a sweet new year.
*Farms with apple orchards may open them to the public, so consumers may themselves pick the apples they will buy.


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 ] began and allowed the development of the multibillion-dollar fruit industry, of which the apple is the leading product.<ref name="UofGeorgia"/>
Sliced apples turn brown with exposure to air due to the conversion of natural phenolic substances into ] upon exposure to ]. Different cultivars differ in their propensity to brown after slicing. Sliced fruit can be treated with ] to prevent this effect.


Until the 20th century, farmers stored apples in ] 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> ] 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=] |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>
==Health benefits==
{{nutritionalvalue | name=Apples, with skin (edible parts) | kJ=218 | protein=0.26 g | fat=0.17 g | carbs=13.81 g | fiber=2.4 g | | sugars=10.39 g | iron_mg=0.12 | calcium_mg=6 | magnesium_mg=5 | phosphorus_mg=11 | potassium_mg=107 | zinc_mg=0.04 | vitC_mg=4.6 | pantothenic_mg=0.061 | vitB6_mg=0.041 | folate_ug=3 | thiamin_mg=0.017 | riboflavin_mg=0.026 | niacin_mg=0.091 | right=1 | source_usda=1 }}


=== Breeding ===
An old ] attests to the health benefits of the fruit: "''An apple a day keeps the doctor away.''" Research suggests that apples may reduce the risk of ], ] and ].<ref>, from Stanford comprehensive cancer center.</ref> Like many fruits, apples contain ] as well as a host of other ] compounds, which may reduce the risk of cancer by preventing ] damage. The fiber content, while less than in most other fruits, helps regulate bowel movements and may thus reduce the risk of colon cancer. They may also help with ], ] and controlling ], as they do not have any cholesterol, have fibre (which reduces cholesterol by preventing reabsorption), and are bulky for their caloric content like most fruits and vegetables.
{{See also|Fruit tree propagation|Malling series}}


]
A group of chemicals in apples could protect the brain from the type of damage that triggers such ]s as ] and ]. Chang Y. 'Cy' Lee of the ] found that the apple ]s, which are naturally occurring ] found in fresh apples, can protect ]s from neurotoxicity induced by ]. The researchers used red delicious apples from New York State to provide the extracts to study the effects of ]. Lee said that all apples are high in the critical ]s and that the amount of phenolic compounds in the apple flesh and in the skin vary from year to year, season to season and from growing region to growing region (November/December 2004 issue of the Journal of Food Science). The predominant phenolic phytochemicals in apples are ], ], and ] B2 (PMID 14558772).


Many apples grow readily from seeds. However, apples must be propagated asexually to obtain cuttings with the characteristics of the parent. This is because seedling apples are "]". 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=] |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> ] 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) |url-status=dead}}</ref>
The ]s are mildly poisonous, containing a small amount of ], a ] ], but a large amount would need to be chewed to have any toxic effect.<ref>{{cite web | url=http://web.archive.org/web/20070613110058/http://rawveg.info/rawfoodtoxins.html | title=Raw Food Toxins | RawVeg.info | accessdate=2006-09-16}}</ref>


Because apples are not ] when planted as seeds, propagation usually involves ] of cuttings. The ] 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 |url-status=dead |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=]}}</ref>
Apple consumption can help remove trapped food and clean between the teeth, but the ] contained within the fruit is also capable of eroding ] over time, and through excess consumption.


Dwarf rootstocks for apple trees can be traced as far back as 300 BCE, to the area of ] and ]. ] sent samples of dwarf apple trees to ]'s ]. 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 ] 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 ']' in ].<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 |url-status=dead}}</ref>
==Cultural aspects==
:''For the allegorical use of the apple in ], ] and ], as well as other cultural aspects, see : ].''


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=978-0851995922 |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 ]s (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=] |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 |url-status=dead}}</ref>
Apples appear in many ], often as a mystical and ]. One of the problems identifying apples in religion, mythology and folktales is that the word "apple" was used as a generic term for all fruit, other than berries but including nuts, as late as the ].


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>
==Storage==
Commercially, apples can be stored for some months in controlled-atmosphere chambers to delay ]-induced onset of ripening. Ripening begins when the fruit is removed.<ref></ref>
For home storage, most varieties of apple can be stored for approximately two weeks, when kept at the coolest part of the refrigerator. Some types of apple, including the ] and ], have an even longer shelf life.<ref> {{cite web | accessdate = 2007-05-25 | url = http://www.foodscience.csiro.au/refrigerated.htm | title = Food Science Australia Fact Sheet: Refrigerated storage of perishable foods | date = February, 2005 | language = English | last = Food Science Australia }}</ref>


==See also== === Pollination ===
{{See also|Fruit tree pollination}}
<div style="-moz-column-count:2; column-count:2;">
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
* ]
</div>


] cultivar]]
==References==
<div class="references-small">
<references/>
</div>


] on an apple bloom in ], Canada]]
==External links ==

{{Wiktionary}} {{Commons|Apple}} {{Cookbook|Apple}}
Apples are self-incompatible; they must ] to develop fruit. During the flowering each season, apple growers often utilize ]s to carry pollen. ]s are most commonly used. ] are also used as supplemental pollinators in commercial orchards. ] ] 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=] |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>
* with some years and places of cultivar origins

* from the UK's ]
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:
* from Midwest Apple Improvement Association

*
* Group A – Early flowering, 1 to 3 May in England (']', 'Red Astrachan')
*
* Group B – 4 to 7 May (']', ']')
* Group C – Mid-season flowering, 8 to 11 May (']', ']')
* Group D – Mid/late season flowering, 12 to 15 May (']', 'Calville blanc d'hiver')
* Group E – Late flowering, 16 to 18 May (']', '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>{{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 |language=en |edition= |location=Wageningen, Netherlands |publisher=] |issue=179 |pages=129–140 |doi=10.17660/ActaHortic.1986.179.10 |isbn=978-90-6605-182-9}}</ref>

=== Maturation and harvest ===
{{See also|Fruit picking|Fruit tree pruning}}
], the former ], with his family picking apples in the 1930s]]

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=live}}</ref>

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

=== Storage ===

] in a wholesale food market]]

Commercially, apples can be stored for months in ] chambers. Apples are commonly stored in chambers with lowered concentrations of ] to reduce respiration and slow softening and other changes if the fruit is already fully ripe. The gas ] is used by plants as a ] 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 ], 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 culitvars of apple can be stored for three weeks in a pantry and four to six weeks from the date of purchase in a refrigerator that maintains {{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=] |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=] |access-date=17 September 2024 |language=en |date=12 April 2019}}</ref> Some varieties of apples (e.g. ']' and ']') 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=] |url-status=dead |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 ] 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=] |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 ===
{{further|List of apple diseases}}

] larva tunnelling inside an apple]]

Apple trees are susceptible to ] and ] diseases, and to damage by insect pests. Many commercial orchards pursue a program of chemical sprays to maintain high fruit quality, tree health, and high yields. These prohibit the use of synthetic pesticides, though some older pesticides are allowed. ] methods include, for instance, introducing its natural predator to reduce the population of a particular pest.

A wide range of pests and diseases can affect the plant. Three of the more common diseases or pests are mildew, aphids, and apple scab.

* ] 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 ]—eliminating the conditions that caused the disease and burning the infected plants are among recommended actions.<ref>{{cite book |last1=Jackson |first1=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 |location=North Yakima, Washington |pages=475–476 |chapter-url=https://archive.org/details/CUbiodiversity628875-8748/page/475 |access-date=1 August 2024 |volume=I |chapter=Powdery Mildew}}</ref>
* ]s are small insects with ]. 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 |language=en |publisher=] |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=bot: unknown }}</ref>
* ]: 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 |language=en |location= New York |publisher=] |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 ], and three fungal diseases: '']'' rust, ],<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=] |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=bot: unknown }}</ref> and ].<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=] |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}}</ref> ]s, and the ]s 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 ] 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}}

]

There are more than 7,500 known ]s (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=] |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 ] and the ultimate size of the tree, even when grown on the same ].<ref name="England" /> Different cultivars are available for ] and ] climates. The UK's National Fruit Collection, which is the responsibility of the Department of Environment, Food, and Rural Affairs, includes a collection of over 2,000 cultivars of apple tree in Kent.<ref>{{cite web |url=http://www.nationalfruitcollection.org.uk/index.php |title=Home|website=] |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 ], which is responsible for developing the UK national collection database, provides access to search the national collection. The University of Reading's work is part of the European Cooperative Programme for Plant Genetic Resources of which there are 38 countries participating in the Malus/Pyrus work group.<ref>{{cite web |url=http://www.ecpgr.cgiar.org/index.php?id=2501&tx_wfqbe_pi1&#91;uid&#93;=59 |title=ECPGR Malus/Pyrus Working Group Members |website=Ecpgr.cgiar.org |date=22 July 2002 |access-date=25 August 2014 |url-status=dead |archive-url=https://web.archive.org/web/20140826114504/http://www.ecpgr.cgiar.org/index.php?id=2501&tx_wfqbe_pi1&#91;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 (]s) or producing ]. ]s 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 |url-status=dead |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 the United States there are many apple breeding programs associated with universities. ] has had a program operating since 1880 in ]. Among their recent well known apples is the 'SnapDragon' cultivar released in 2013. In the west ] started a program to support their apple industry in 1994 and released the ']' cultivar in 2017. The third most grown apple cultivar in the United States is the ']', released by the ] program in 1991.<ref>{{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> In Europe there are also many breeding programs such as the ], 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. |title=Resistance Breeding in Apple at Dresden-Pillnitz |journal=Ecofruit - 13th International Conference on Cultivation Technique and Phytopathological Problems in Organic Fruit-Growing: Proceedings to the Conference from 18thFebruary to 20th February 2008 at Weinsberg/Germany |date=2008 |pages=220–225 |url=http://orgprints.org/13698/ |access-date=31 July 2024 |language=de |archive-date=28 January 2021 |archive-url=https://web.archive.org/web/20210128124933/https://orgprints.org/13698/ |url-status=live }}</ref>

Commercially popular apple cultivars are soft but crisp. Other desirable qualities in modern commercial apple breeding are a colorful skin, absence of ], 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=] |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. Most North Americans and Europeans favor sweet, subacid apples, but tart apples have a strong minority following.<ref name="World" /> 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 |url-status=dead }}</ref> especially the ].<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 that sell directly to local markets. Many unusual and locally important cultivars with their own unique taste and appearance exist; apple conservation campaigns have sprung up around the world to preserve such local cultivars from extinction. In the United Kingdom, old cultivars such as ']' and ']' 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=] |archive-url=https://web.archive.org/web/20080121045236/http://www.uga.edu/fruit/apple.html |archive-date=21 January 2008 |url-status=dead}}</ref>
<!--
Please do not add a gallery of cultivars here, that job is already done at ], which is "main" linked at the top of this section. Thanks.
-->

=== Production ===
{{Main|List of countries by apple production}}

{| class="wikitable floatright" style="width:13em; text-align:center"
! colspan=2|Apple production<br>
<small>2022, millions of ]s</small><br/>
|-
|{{CHN}}|| 47.6
|-
|{{USA}}|| 4.8
|-
|{{TUR}}|| 4.4
|-
|{{POL}}|| 4.3
|-
|{{IND}}|| 2.6
|-
|'''World''' || '''95.8'''
|-
|colspan=2 style="text-align: center;" | <small>Source: ] of the United Nations</small><ref name="faostat">{{cite web |date=2024 |title=Apple production in 2022; from pick lists: Crops/World Regions/Production Quantity |url=http://www.fao.org/faostat/en/#data/QC |access-date=18 June 2024 |publisher=FAOSTAT, UN ], Statistics Division |archive-date=12 November 2016 |archive-url=https://web.archive.org/web/20161112130804/http://www.fao.org/faostat/en/#data/QC |url-status=live }}</ref>
|}

World production of apples in 2022 was 96 million ]s, with China producing 50% of the total (table).<ref name=faostat/> Secondary producers were the United States, ], and ].<ref name=faostat/>

== Toxicity ==

=== Amygdalin ===
Apple seeds contain small amounts of ], a sugar and ] compound known as a ]. Ingesting small amounts of apple seeds causes no ill effects, but consumption of extremely large doses can cause ]s. It may take several hours before the poison takes effect, as cyanogenic glycosides must be ] 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=] : ] |pages=27, 211–212 |isbn=978-0387-31268-2 |lccn=2005938815 |oclc=77537459 |access-date=11 September 2024}}</ref> The U.S. ]'s ] 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 |url-status=live |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 ] ].<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 ] (OAS), generally involve itching and inflammation of the mouth and throat,<ref name="EU" /> but in rare cases can also include life-threatening ].<ref>Landau, Elizabeth, , 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 ], 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 ] fruit suitable for either of the two forms of apple allergy.<ref name="EU"/>

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

=== Nutrition ===
{{stack|
{{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
|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 =
}}
}}

A raw apple is 86% water and 14% ]s, with negligible content of ] and ] (table). A reference serving of a raw apple with skin weighing {{cvt|100|g}} provides 52 ]s and a moderate content of ] (table). Otherwise, there is low content of ]s, with the ]s of all falling below 10% (table).

=== Culinary ===

{{further|List of apple dishes}}

]

Apples varieties can be grouped as ]s, ], and ]s, the last so astringent as to be "almost inedible".<ref name="Davidson 2014"/> Apples are consumed as ], raw in salads, baked in ], cooked into ] and ], 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=] |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 ].<ref name="Davidson 2014"/> Juice and cider are produced commercially; cider is a significant industry in regions such as the ] and ].<ref name="Davidson 2014"/>

A ] (UK) or ] (US) is a confection made by coating an apple in hot ] or ] 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=] |language=en |edition=Third |location=Oxford |publisher=] |page=824 |isbn=978-0-19-967733-7 |lccn=2013957569 |oclc=890807357 |ol=27172691M |access-date=18 September 2024}}</ref><ref name="Lim 2012" /> ] are a ritual ] eaten during the Jewish New Year of ].<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 ], ]s, and ]. When cooked, some apple cultivars easily form a puree known as ], which can be cooked down to form a preserve, apple butter. They are often ] or ], 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=] |language=en |edition=Third |location=Oxford |publisher=] |pages=27–31 |isbn=978-0-19-967733-7 |lccn=2013957569 |oclc=890807357 |ol=27172691M |access-date=18 September 2024}}</ref>

{{Cookbook}}
Apples are ] or ] to produce ], which may be drunk unfiltered (called ] in North America), or filtered. Filtered juice is often concentrated and frozen, then reconstituted later and consumed. Apple juice can be ] to make ] (called hard cider in North America), ], and vinegar.<ref name="Lim 2012" /> Through ], various alcoholic beverages can be produced, such as ], ], and ].<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===
] 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 |url-status=dead |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 (], ]) has been approved by the EU Organic Standing Committee to improve organic yield and quality.<ref name=croplife/> A light coating of ], 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 ]s.<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 ] ], which causes ] in sliced or bruised apples, by ] the ] of phenolic compounds to ]s, 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. ], a non-browning group of apples introduced to the United States market in 2019, have been ] to silence the ] of polyphenol oxidase, thereby delaying a browning effect and improving apple eating quality.<ref>{{cite web |title=PPO silencing |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 ] in 2015, and ] 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 |url-status=live |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 ===
] is obtained by ] apple seeds for manufacturing ].<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 ===
]" (1901) by ]]]

In ], the goddess ] is portrayed in the '']'' (written in the 13th century by ]) as providing apples to the ] that give them ]fulness. The English scholar ] links apples to religious practices in ], from which ] developed. She points out that buckets of apples were found in the ] burial site in Norway, that fruit and nuts (Iðunn having been described as being transformed into a nut in '']'') have been found in the early graves of the ] in England and elsewhere on the continent of Europe, which may have had a symbolic meaning, and that nuts are still a recognized symbol of ] 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=] |pages=165–166 |isbn=0-14-013627-4 |oclc=29336401}}</ref>

Davidson notes a connection between apples and the ], a tribe of gods associated with ] in Norse mythology, citing an instance of eleven "golden apples" being given to woo the beautiful ] by ], who was acting as messenger for the major Vanir god ] in stanzas 19 and 20 of '']''. Davidson also notes a further connection between fertility and apples in Norse mythology in chapter 2 of the '']'': when the major goddess ] sends King ] 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 ].<ref name="Davidson 1990" /> Rerir's wife's consumption of the apple results in a six-year pregnancy and the birth (by ]) of their son—the hero ].<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=] |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 ]" used in an 11th-century poem by the ] 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 ] 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 ] and came to Europe from the ], 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 ===
] with the apple of ] ]]

Apples appear in many ], including Greek and Roman ] 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 ], the ] ], as a part of his ], was required to travel to the Garden of the Hesperides and pick the golden apples off the ] 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=] |pages=64–70 |isbn=978-0-89089-924-3 |lccn=00040351 |oclc=46337324}}</ref>

The Greek goddess of discord, ], became disgruntled after she was excluded from the wedding of ] and ].<ref>{{cite web |title=Eris - Greek Goddess of Strife & Discord (Roman Discordia) |url=https://www.theoi.com/Daimon/Eris.html |website=] |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 ] inscribed ] (''Kallistē'', "For the most beautiful one"), into the wedding party. Three goddesses claimed the apple: ], ], and ]. ] of ] 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, ] of ]. He awarded the apple to Aphrodite, thus indirectly causing the ].<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=] |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=] |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=] |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 = ]
| source = Epigram VII
}}

], also of Greek mythology, raced all her suitors in an attempt to avoid marriage. She outran all but ] (also known as ], 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=] |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 ], the ] has many names, including ''Emain Ablach'', "Emain of the Apple-trees". A version of this is ] in ], or in ] ''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=] |pages=122–123 |isbn=978-0-87338-824-5 |lccn=2004024490 |oclc=56805947}}</ref>

=== China ===
]

In China, apples symbolise ], since the sounds of the first element ("píng") in the words "apple" (苹果, ''Píngguǒ'') and "peace" (平安, ''Píng'ān'') are ] 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 ] in Mandarin is ''Píngānyè'' (平安夜, "Peaceful/Quiet Evening"), which made the ]ing 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 ===
]'' by ] (1507), showcasing the apple as a symbol of sin]]

Though the ] of ] in the ] is not identified, popular Christian tradition has held that it was an apple that ] coaxed ] to share with her.<ref name="Macrone">{{cite book |last1=Macrone |first1=Michael |title=Brush up your Bible! |date=1998 |publisher=] |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 ] 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=] |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>

] painters may also have been influenced by the story of the ]s in the ]. 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 ] in the human throat has been called the "]" because of a notion that it was caused by the forbidden fruit remaining in the throat of Adam. The apple as symbol of sexual ] has been used to imply human sexuality, possibly in an ironic vein.<ref name="Macrone"/>

=== Proverb ===
The ], "]", addressing the supposed health benefits of the fruit, has been traced to 19th-century ], 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=] |pages=9, 22, 50 |isbn=978-0-375-50129-6 |lccn=00066479 |oclc=49803415}}</ref>

== See also ==

* ]
* ], apple–crabapple hybrids for eating
* ]

== 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=] |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=] |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=] |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=] |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=] |location=London |isbn=9780711231412 |edition=Second |language=en |oclc=646397065 |ref=none}}

== External links ==

* {{Commons category-inline|Apples}}


{{Apples}} {{Apples}}
{{Crabapple}}
{{Taxonbar|from=Q18674606|from2=Q158657}}
{{Authority control}}


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Latest revision as of 23:48, 20 December 2024

Fruit that grows on a tree This article is about the fruit. For the technology company, see Apple Inc. For other uses, see Apple (disambiguation). "Apple tree" redirects here. For other uses, see Apple tree (disambiguation).

Apple
'Cripps Pink' apples
Flowers
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Rosales
Family: Rosaceae
Genus: Malus
Species: M. domestica
Binomial name
Malus domestica
(Suckow) Borkh.
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

An apple is a round, edible fruit produced by an apple tree (Malus spp., among them the domestic or orchard apple; Malus domestica). Apple trees are cultivated worldwide and are the most widely grown species in the genus Malus. The tree originated in Central Asia, where its wild ancestor, Malus sieversii, is still found. Apples have been grown for thousands of years in Eurasia and were introduced to North America by European colonists. Apples have religious and mythological significance in many cultures, including Norse, Greek, and European Christian tradition.

Apples grown from seed tend to be very different from those of their parents, and the resultant fruit frequently lacks desired characteristics. For commercial purposes, including botanical evaluation, apple cultivars are propagated by clonal grafting onto rootstocks. Apple trees grown without rootstocks tend to be larger and much slower to fruit after planting. Rootstocks are used to control the speed of growth and the size of the resulting tree, allowing for easier harvesting.

There are more than 7,500 cultivars of apples. Different cultivars are bred for various tastes and uses, including cooking, eating raw, and cider or apple juice production. Trees and fruit are prone to fungal, bacterial, and pest problems, which can be controlled by a number of organic and non-organic means. In 2010, the fruit's genome was sequenced as part of research on disease control and selective breeding in apple production.

Etymology

The word apple, whose Old English ancestor is æppel, is descended from the Proto-Germanic noun *aplaz, descended in turn from Proto-Indo-European *h₂ébōl. As late as the 17th century, the word also functioned as a generic term for all fruit, including nuts. This can be compared to the 14th-century Middle English expression appel of paradis, meaning a banana.

Description

The apple is a deciduous tree, generally standing 2 to 4.5 metres (6 to 15 feet) tall in cultivation and up to 15 m (49 ft) in the wild, though more typically 2 to 10 m (6.5 to 33 ft). When cultivated, the size, shape and branch density are determined by rootstock selection and trimming method. Apple trees may naturally have a rounded to erect crown with a dense canopy of leaves. The bark of the trunk is dark gray or gray-brown, but young branches are reddish or dark-brown with a smooth texture. Young twigs are covered in fine downy hairs; they become hairless when older.

The buds are egg-shaped and dark red or purple in color; they range in size from 3 to 5 millimeters, but are usually less than 4 mm. The bud scales have very hairy edges. When emerging from the buds, the leaves are convolute, meaning that their edges overlap each other. Leaves can be simple ovals (elliptic), medium or wide in width, somewhat egg-shaped with the wider portion toward their base (ovate), or even with sides that are more parallel to each other instead of curved (oblong) with a narrow pointed end. The edges have broadly-angled teeth, but do not have lobes. The top surface of the leaves are glabrescent, almost hairless, while the undersides are densely covered in fine hairs. The leaves are attached alternately by short leaf stems 1-to-3.5 cm (1⁄2-to-1+1⁄2 in) long.

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

  • Apple blossoms Apple blossoms
  • Botanical illustration Botanical illustration

Fruit

The fruit is a pome that matures in late summer or autumn. The true fruits or carpels are the harder interior chambers inside the apple's core. There are usually five carpels inside an apple, but there may be as few as three. Each of the chambers contains one or two seeds. The edible flesh is formed from the receptacle at the base of the flower.

  • How apple fruit derives from flower structures 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. The five sepals remain attached and stand out from the surface of the apple.

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

The groundcolor of ripe apples is yellow, green, yellow-green or whitish yellow. The overcolor of ripe apples can be orange-red, pink-red, red, purple-red or brown-red. The overcolor amount can be 0–100%. The skin may be wholly or partly russeted, making it rough and brown. The skin is covered in a protective layer of epicuticular wax. The skin may also be marked with scattered dots. The flesh is generally pale yellowish-white, though it can be pink, yellow or green.

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

Chemistry

Important volatile compounds in apples that contribute to their scent and flavour include acetaldehyde, ethyl acetate, 1-butanal, ethanol, 2-methylbutanal, 3-methylbutanal, ethyl propionate, ethyl 2-methylpropionate, ethyl butyrate, ethyl 2-methyl butyrate, hexanal, 1-butanol, 3-methylbutyl acetate, 2-methylbutyl acetate, 1-propyl butyrate, ethyl pentanoate, amyl acetate, 2-methyl-1-butanol, trans-2-hexenal, ethyl hexanoate, hexanol.

Taxonomy

The apple as a species has more than 100 alternative scientific names, or synonyms. In modern times, Malus pumila and Malus domestica are the two main names in use. M. pumila is the older name, but M. domestica has become much more commonly used starting in the 21st century, especially in the western world. Two proposals were made to make M. domestica a conserved name: the earlier proposal was voted down by the Committee for Vascular Plants of the IAPT in 2014, but in April 2017 the Committee decided, with a narrow majority, that the newly popular name should be conserved. The General Committee of the IAPT decided in June 2017 to approve this change, officially conserving M. domestica. Nevertheless, some works published after 2017 still use M. pumila as the correct name, under an alternate taxonomy.

When first classified by Linnaeus in 1753, the pears, apples, and quinces were combined into one genus that he named Pyrus and he named the apple as Pyrus malus. This was widely accepted, however the botanist Philip Miller published an alternate classification in The Gardeners Dictionary with the apple species separated from Pyrus in 1754. He did not clearly indicate that by Malus pumila he meant the domesticated apple. Nonetheless, it was used as such by many botanists. When Moritz Balthasar Borkhausen published his scientific description of the apple in 1803 it may have been a new combination of P. malus var. domestica, but this was not directly referenced by Borkhausen. The earliest use of var. domestica for the apple was by Georg Adolf Suckow in 1786.

Genome

Further information: 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'. However, this first whole genome sequence contained several errors, 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.

The first whole genome assembly was estimated to contain around 57,000 genes, though the more recent genome sequences support estimates between 42,000 and 44,700 protein-coding genes. The availability of whole genome sequences has provided evidence that the wild ancestor of the cultivated apple most likely is Malus sieversii. Re-sequencing of multiple accessions has supported this, while also suggesting extensive introgression from Malus sylvestris following domestication.

Cultivation

History

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.
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. 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. Cultivation of the species, most likely beginning on the forested flanks of the Tian Shan mountains, progressed over a long period of time and permitted secondary introgression of genes from other species into the open-pollinated seeds. Significant exchange with Malus sylvestris, the crabapple, resulted in populations of apples being more related to crabapples than to the more morphologically similar progenitor Malus sieversii. In strains without recent admixture the contribution of the latter predominates.

The apple is thought to have been domesticated 4,000–10,000 years ago in the Tian Shan mountains, and then to have travelled along the Silk Road to Europe, with hybridization and introgression of wild crabapples from Siberia (M. baccata), the Caucasus (M. orientalis), and Europe (M. sylvestris). Only the M. sieversii trees growing on the western side of the Tian Shan mountains contributed genetically to the domesticated apple, not the isolated population on the eastern side.

Chinese soft apples, such as M. asiatica and M. prunifolia, have been cultivated as dessert apples for more than 2,000 years in China. These are thought to be hybrids between M. baccata and M. sieversii in Kazakhstan.

Among the traits selected for by human growers are size, fruit acidity, color, firmness, and soluble sugar. Unusually for domesticated fruits, the wild M. sieversii origin is only slightly smaller than the modern domesticated apple.

At the Sammardenchia-Cueis site near Udine in Northeastern Italy, seeds from some form of apples have been found in material carbon dated to between 6570 and 5684 BCE. Genetic analysis has not yet been successfully used to determine whether such ancient apples were wild Malus sylvestris or Malus domesticus containing Malus sieversii ancestry. It is hard to distinguish in the archeological record between foraged wild apples and apple plantations.

There is indirect evidence of apple cultivation in the third millennium BCE in the Middle East. There is direct evidence, apple cores, dated to the 10th century BCE from a Judean site between the Sinai and Negev. There was substantial apple production in European classical antiquity, and grafting was certainly known then. Grafting is an essential part of modern domesticated apple production, to be able to propagate the best cultivars; it is unclear when apple tree grafting was invented.

"Wild Apples"
by Henry David Thoreau
Read by Kevin S for LibriVox
Audio 01:01:35 (full text)
Problems playing this file? See media help.

The Roman writer Pliny the Elder describes a method of storage for apples from his time in the 1st century. He says they should be placed in a room with good air circulation from a north facing window on a bed of straw, chaff, or mats with windfalls kept separately. Though methods like this will extend the availabity of reasonably fresh apples, without refrigeration their lifespan is limited. Even sturdy winter apple varieties will only keep well until December in cool climates. For longer storage medieval Europeans strung up cored and peeled apples to dry, either whole or sliced into rings.

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

Apple cultivars brought as seed from Europe were spread along Native American trade routes, as well as being cultivated on colonial farms. An 1845 United States apples nursery catalogue sold 350 of the "best" cultivars, showing the proliferation of new North American cultivars by the early 19th century. In the 20th century, irrigation projects in Eastern Washington began and allowed the development of the multibillion-dollar fruit industry, of which the apple is the leading product.

Until the 20th century, farmers stored apples in frostproof cellars during the winter for their own use or for sale. Improved transportation of fresh apples by train and road replaced the necessity for storage. Controlled atmosphere facilities are used to keep apples fresh year-round. Controlled atmosphere facilities use high humidity, low oxygen, and controlled carbon dioxide levels to maintain fruit freshness. They were first researched at Cambridge University in the 1920s and first used in the United States in the 1950s.

Breeding

See also: Fruit tree propagation and Malling series
An apple tree in Germany

Many apples grow readily from seeds. However, apples must be propagated asexually to obtain cuttings with the characteristics of the parent. This is because seedling apples are "extreme heterozygotes". Rather than resembling their parents, seedlings are all different from each other and from their parents. Triploid cultivars have an additional reproductive barrier in that three sets of chromosomes cannot be divided evenly during meiosis, yielding unequal segregation of the chromosomes (aneuploids). Even in the case when a triploid plant can produce a seed (apples are an example), it occurs infrequently, and seedlings rarely survive.

Because apples are not true breeders when planted as seeds, propagation usually involves grafting of cuttings. The rootstock used for the bottom of the graft can be selected to produce trees of a large variety of sizes, as well as changing the winter hardiness, insect and disease resistance, and soil preference of the resulting tree. Dwarf rootstocks can be used to produce very small trees (less than 3.0 m or 10 ft high at maturity), which bear fruit many years earlier in their life cycle than full size trees, and are easier to harvest.

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. The majority of the rootstocks used to control size in apples were developed in England in the early 1900s. The East Malling Research Station conducted extensive research into rootstocks, and their rootstocks are given an "M" prefix to designate their origin. Rootstocks marked with an "MM" prefix are Malling-series cultivars later crossed with trees of 'Northern Spy' in Merton, England.

Most new apple cultivars originate as seedlings, which either arise by chance or are bred by deliberately crossing cultivars with promising characteristics. The words "seedling", "pippin", and "kernel" in the name of an apple cultivar suggest that it originated as a seedling. Apples can also form bud sports (mutations on a single branch). Some bud sports turn out to be improved strains of the parent cultivar. Some differ sufficiently from the parent tree to be considered new cultivars.

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

Pollination

See also: Fruit tree pollination
Apple blossom from an old Ayrshire cultivar
An orchard mason bee on an apple bloom in British Columbia, Canada

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.

Cultivars are sometimes classified by the day of peak bloom in the average 30-day blossom period, with pollinizers selected from cultivars within a 6-day overlap period. There are four to seven pollination groups in apples, depending on climate:

  • Group A – Early flowering, 1 to 3 May in England ('Gravenstein', 'Red Astrachan')
  • Group B – 4 to 7 May ('Idared', 'McIntosh')
  • Group C – Mid-season flowering, 8 to 11 May ('Granny Smith', 'Cox's Orange Pippin')
  • Group D – Mid/late season flowering, 12 to 15 May ('Golden Delicious', 'Calville blanc d'hiver')
  • Group E – Late flowering, 16 to 18 May ('Braeburn', 'Reinette d'Orléans')
  • Group F – 19 to 23 May ('Suntan')
  • Group H – 24 to 28 May ('Court-Pendu Gris' – also called Court-Pendu plat)

One cultivar can be pollinated by a compatible cultivar from the same group or close (A with A, or A with B, but not A with C or D).

Maturation and harvest

See also: Fruit picking and Fruit tree pruning
L. K. Relander, the former President of Finland, with his family picking apples in the 1930s

Cultivars vary in their yield and the ultimate size of the tree, even when grown on the same rootstock. Some cultivars, if left unpruned, grow very large—letting them bear more fruit, but making harvesting more difficult. Depending on tree density (number of trees planted per unit surface area), mature trees typically bear 40–200 kg (90–440 lb) of apples each year, though productivity can be close to zero in poor years. Apples are harvested using three-point ladders that are designed to fit amongst the branches. Trees grafted on dwarfing rootstocks bear about 10–80 kg (20–180 lb) of fruit per year.

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

Crops ripen at different times of the year according to the cultivar. Cultivar that yield their crop in the summer include 'Sweet Bough' and 'Duchess'; fall producers include 'Blenheim'; winter producers include 'King', 'Swayzie', and 'Tolman Sweet'.

Storage

Different apple cultivars in a wholesale food market

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.

For home storage, most culitvars of apple can be stored for three weeks in a pantry and four to six weeks from the date of purchase in a refrigerator that maintains 4 to 0 °C (39 to 32 °F). Some varieties of apples (e.g. 'Granny Smith' and 'Fuji') have more than three times the storage life of others.

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

Pests and diseases

Further information: List of apple diseases
Codling moth larva tunnelling inside an apple

Apple trees are susceptible to fungal and bacterial diseases, and to damage by insect pests. Many commercial orchards pursue a program of chemical sprays to maintain high fruit quality, tree health, and high yields. These prohibit the use of synthetic pesticides, though some older pesticides are allowed. Organic methods include, for instance, introducing its natural predator to reduce the population of a particular pest.

A wide range of pests and diseases can affect the plant. Three of the more common diseases or pests are mildew, aphids, and apple scab.

  • Mildew is characterized by light grey powdery patches appearing on the leaves, shoots and flowers, normally in spring. The flowers turn a creamy yellow color and do not develop correctly. This can be treated similarly to Botrytis—eliminating the conditions that caused the disease and burning the infected plants are among recommended actions.
  • Aphids are small insects with sucking mouthparts. Five species of aphids commonly attack apples: apple grain aphid, rosy apple aphid, apple aphid, spirea aphid, and the woolly apple aphid. The aphid species can be identified by color, time of year, and by differences in the cornicles (small paired projections from their rear). Aphids feed on foliage using needle-like mouth parts to suck out plant juices. When present in high numbers, certain species reduce tree growth and vigor.
  • Apple scab: Apple scab causes leaves to develop olive-brown spots with a velvety texture that later turn brown and become cork-like in texture. The disease also affects the fruit, which also develops similar brown spots with velvety or cork-like textures. Apple scab is spread through fungus growing in old apple leaves on the ground and spreads during warm spring weather to infect the new year's growth.

Among the most serious disease problems is a bacterial disease called fireblight, and three fungal diseases: Gymnosporangium rust, black spot, and bitter rot. Codling moths, and the apple maggots of fruit flies, cause serious damage to apple fruits, making them unsaleable. Young apple trees are also prone to mammal pests like mice and deer, which feed on the soft bark of the trees, especially in winter. The larvae of the apple clearwing moth (red-belted clearwing) burrow through the bark and into the phloem of apple trees, potentially causing significant damage.

Cultivars

Main article: List of apple cultivars
An assortment of apple cultivars

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

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

In the United States there are many apple breeding programs associated with universities. Cornell University has had a program operating since 1880 in Geneva, New York. Among their recent well known apples is the 'SnapDragon' cultivar released in 2013. In the west Washington State University started a program to support their apple industry in 1994 and released the 'Cosmic Crisp' cultivar in 2017. The third most grown apple cultivar in the United States is the 'Honeycrisp', released by the University of Minnesota program in 1991. Unusually for a popular cultivar, the 'Honeycrisp' is not directly related to another popular apple cultivar but instead to two unsuccessful cultivars. In Europe there are also many breeding programs such as the Julius Kühn-Institut, the German federal research center for cultivated plants.

Commercially popular apple cultivars are soft but crisp. Other desirable qualities in modern commercial apple breeding are a colorful skin, absence of russeting, ease of shipping, lengthy storage ability, high yields, disease resistance, common apple shape, and developed flavor. Modern apples are generally sweeter than older cultivars, as popular tastes in apples have varied over time. Most North Americans and Europeans favor sweet, subacid apples, but tart apples have a strong minority following. Extremely sweet apples with barely any acid flavor are popular in Asia, especially the Indian subcontinent.

Less common apple cultivars from an orchard in Italy

Old cultivars are often oddly shaped, russeted, and grow in a variety of textures and colors. Some find them to have better flavor than modern cultivars, but they may have other problems that make them commercially unviable—low yield, disease susceptibility, poor tolerance for storage or transport, or just being the "wrong" size. A few old cultivars are still produced on a large scale, but many have been preserved by home gardeners and farmers that sell directly to local markets. Many unusual and locally important cultivars with their own unique taste and appearance exist; apple conservation campaigns have sprung up around the world to preserve such local cultivars from extinction. In the United Kingdom, old cultivars such as 'Cox's Orange Pippin' and 'Egremont Russet' are still commercially important even though by modern standards they are low yielding and susceptible to disease.

Production

Main article: List of countries by apple production
Apple production

2022, millions of tonnes

 China 47.6
 United States 4.8
 Turkey 4.4
 Poland 4.3
 India 2.6
World 95.8
Source: FAOSTAT of the United Nations

World production of apples in 2022 was 96 million tonnes, with China producing 50% of the total (table). Secondary producers were the United States, Turkey, and Poland.

Toxicity

Amygdalin

Apple seeds contain small amounts of amygdalin, a sugar and cyanide compound known as a cyanogenic glycoside. Ingesting small amounts of apple seeds causes no ill effects, but consumption of extremely large doses can cause adverse reactions. It may take several hours before the poison takes effect, as cyanogenic glycosides must be hydrolyzed before the cyanide ion is released. The U.S. National Library of Medicine's Hazardous Substances Data Bank records no cases of amygdalin poisoning from consuming apple seeds.

Allergy

One form of apple allergy, often found in northern Europe, is called birch-apple syndrome and is found in people who are also allergic to birch pollen. Allergic reactions are triggered by a protein in apples that is similar to birch pollen, and people affected by this protein can also develop allergies to other fruits, nuts, and vegetables. Reactions, which entail oral allergy syndrome (OAS), generally involve itching and inflammation of the mouth and throat, but in rare cases can also include life-threatening anaphylaxis. This reaction only occurs when raw fruit is consumed—the allergen is neutralized in the cooking process. The variety of apple, maturity and storage conditions can change the amount of allergen present in individual fruits. Long storage times can increase the amount of proteins that cause birch-apple syndrome.

In other areas, such as the Mediterranean, some individuals have adverse reactions to apples because of their similarity to peaches. This form of apple allergy also includes OAS, but often has more severe symptoms, such as vomiting, abdominal pain and urticaria, and can be life-threatening. Individuals with this form of allergy can also develop reactions to other fruits and nuts. Cooking does not break down the protein causing this particular reaction, so affected individuals cannot eat raw or cooked apples. Freshly harvested, over-ripe fruits tend to have the highest levels of the protein that causes this reaction.

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

Uses

See also: Cooking apple and Cider apple

Nutrition

Apples, with skin (edible parts)
Nutritional value per 100 g (3.5 oz)
Energy218 kJ (52 kcal)
Carbohydrates13.81 g
Sugars10.39
Dietary fiber2.4 g
Fat0.17 g
Protein0.26 g
Vitamins and minerals
VitaminsQuantity %DV
Vitamin A equiv.beta-Carotenelutein zeaxanthin0% 3 μg0%27 μg29 μg
Thiamine (B1)1% 0.017 mg
Riboflavin (B2)2% 0.026 mg
Niacin (B3)1% 0.091 mg
Pantothenic acid (B5)1% 0.061 mg
Vitamin B62% 0.041 mg
Folate (B9)1% 3 μg
Vitamin C5% 4.6 mg
Vitamin E1% 0.18 mg
Vitamin K2% 2.2 μg
MineralsQuantity %DV
Calcium0% 6 mg
Iron1% 0.12 mg
Magnesium1% 5 mg
Manganese2% 0.035 mg
Phosphorus1% 11 mg
Potassium4% 107 mg
Sodium0% 1 mg
Zinc0% 0.04 mg
Other constituentsQuantity
Water85.56 g

Link to Full Nutrient Report of USDA Database entry
Percentages estimated using US recommendations for adults, except for potassium, which is estimated based on expert recommendation from the National Academies.

A raw apple is 86% water and 14% carbohydrates, with negligible content of fat and protein (table). A reference serving of a raw apple with skin weighing 100 g (3.5 oz) provides 52 calories and a moderate content of dietary fiber (table). Otherwise, there is low content of micronutrients, with the Daily Values of all falling below 10% (table).

Culinary

Further information: List of apple dishes
Machine for paring, coring, and slicing apples, from Henry B. Scammell's 1897 handbook Cyclopedia of Valuable Receipts

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

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

A toffee apple (UK) or caramel apple (US) is a confection made by coating an apple in hot toffee or caramel candy respectively and allowing it to cool. Apples and honey are a ritual food pairing eaten during the Jewish New Year of Rosh Hashanah.

Apples are an important ingredient in many desserts, such as pies, crumbles, and cakes. When cooked, some apple cultivars easily form a puree known as apple sauce, which can be cooked down to form a preserve, apple butter. They are often baked or stewed, and are cooked in some meat dishes.

Apples are milled or pressed to produce apple juice, which may be drunk unfiltered (called apple cider in North America), or filtered. Filtered juice is often concentrated and frozen, then reconstituted later and consumed. Apple juice can be fermented to make cider (called hard cider in North America), ciderkin, and vinegar. Through distillation, various alcoholic beverages can be produced, such as applejack, Calvados, and apple brandy.

Organic production

Organic apples are commonly produced in the United States. Due to infestations by key insects and diseases, organic production is difficult in Europe. The use of pesticides containing chemicals, such as sulfur, copper, microorganisms, viruses, clay powders, or plant extracts (pyrethrum, neem) has been approved by the EU Organic Standing Committee to improve organic yield and quality. A light coating of kaolin, which forms a physical barrier to some pests, also may help prevent apple sun scalding.

Non-browning apples

Apple skins and seeds contain polyphenols. 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. Browning reduces apple taste, color, and food value. Arctic apples, a non-browning group of apples introduced to the United States market in 2019, have been genetically modified to silence the expression of polyphenol oxidase, thereby delaying a browning effect and improving apple eating quality. The US Food and Drug Administration in 2015, and Canadian Food Inspection Agency in 2017, determined that Arctic apples are as safe and nutritious as conventional apples.

Other products

Apple seed oil is obtained by pressing apple seeds for manufacturing cosmetics.

In culture

Main article: Apple (symbolism)

Germanic paganism

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, that fruit and nuts (Iðunn having been described as being transformed into a nut in Skáldskaparmál) have been found in the early graves of the Germanic peoples in England and elsewhere on the continent of Europe, which may have had a symbolic meaning, and that nuts are still a recognized symbol of fertility in southwest England.

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

Further, Davidson points out the "strange" phrase "Apples of Hel" used in an 11th-century poem by the skald Thorbiorn Brúnarson. She states this may imply that the apple was thought of by Brúnarson as the food of the dead. Further, Davidson notes that the potentially Germanic goddess Nehalennia is sometimes depicted with apples and that parallels exist in early Irish stories. Davidson asserts that while cultivation of the apple in Northern Europe extends back to at least the time of the Roman Empire and came to Europe from the Near East, the native varieties of apple trees growing in Northern Europe are small and bitter. Davidson concludes that in the figure of Iðunn "we must have a dim reflection of an old symbol: that of the guardian goddess of the life-giving fruit of the other world."

Greek mythology

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. In Greek mythology, the Greek hero Heracles, as a part of his Twelve Labours, was required to travel to the Garden of the Hesperides and pick the golden apples off the Tree of Life growing at its center.

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

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:

I throw the apple at you, and if you are willing to love me, take it and share your girlhood with me; but if your thoughts are what I pray they are not, even then take it, and consider how short-lived is beauty.

— Plato, Epigram VII

Atalanta, also of Greek mythology, raced all her suitors in an attempt to avoid marriage. She outran all but Hippomenes (also known as Melanion, a name possibly derived from melon, the Greek word for both "apple" and fruit in general), who defeated her by cunning, not speed. Hippomenes knew that he could not win in a fair race, so he used three golden apples (gifts of Aphrodite, the goddess of love) to distract Atalanta. It took all three apples and all of his speed, but Hippomenes was finally successful, winning the race and Atalanta's hand.

Celtic mythology

In Celtic mythology, the otherworld has many names, including Emain Ablach, "Emain of the Apple-trees". A version of this is Avalon in Arthurian legend, or in Welsh Ynys Afallon, "Island of Apples".

China

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. When these two words are combined, the word Píngānguǒ (平安果, "Peace apples") is formed. This association developed further as the name for Christmas Eve in Mandarin is Píngānyè (平安夜, "Peaceful/Quiet Evening"), which made the gifting of apples at this season to friends and associates popular, as a way to wish them peace and safety.

Christian art

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. 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. The tree of the forbidden fruit is called "the tree of the knowledge of good and evil" in Genesis 2:17, and the Latin for "good and evil" is bonum et malum.

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

Proverb

The proverb, "An apple a day keeps the doctor away", addressing the supposed health benefits of the fruit, has been traced to 19th-century Wales, where the original phrase was "Eat an apple on going to bed, and you'll keep the doctor from earning his bread". In the 19th century and early 20th, the phrase evolved to "an apple a day, no doctor to pay" and "an apple a day sends the doctor away"; the phrasing now commonly used was first recorded in 1922.

See also

References

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Further reading

External links

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Apples
Species
Table apples
Cooking apples
Cider apples
Ornamental apple
Apple products
Food
Drink
Agriculture
"Crabapple" or "Wild apple" (of the genus Malus)
Species, varieties
and cultivars
Crab apples by the roadside
Topics
Taxon identifiers
Malus domestica
Malus pumila
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