Taxonomy

The apple as a species has more than 100 alternative scientific names, or synonyms.[17] In
modern times, Malus pumila and Malus domestica are the two main names in use. M. pumila is
the older name, but M. domestica has become much more commonly used starting in the 21st
century, especially in the western world. Two proposals were made to
make M. domestica a conserved name: the earlier proposal was voted down by the Committee
for Vascular Plants of the IAPT in 2014, but in April 2017 the Committee decided, with a narrow
majority, that the newly popular name should be conserved.[18] The General Committee of the
IAPT decided in June 2017 to approve this change, officially conserving M. domestica.
[19]
Nevertheless, some works published after 2017 still use M. pumila as the correct name,
under an alternate taxonomy.[2]

When first classified by Linnaeus in 1753, the pears, apples, and quinces were combined into
one genus that he named Pyrus and he named the apple as Pyrus malus. This was widely
accepted, however the botanist Philip Miller published an alternate classification in The
Gardeners Dictionary with the apple species separated from Pyrus in 1754. He did not clearly
indicate that by Malus pumila he meant the domesticated apple. Nonetheless, it was used as
such by many botanists. When Moritz Balthasar Borkhausen published his scientific description
of the apple in 1803 it may have been a new combination of P. malus var. domestica, but this
was not directly referenced by Borkhausen.[17] The earliest use of var. domestica for the apple
was by Georg Adolf Suckow in 1786.[2]

Genome

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'.[20] However,
this first whole genome sequence contained several errors,[21] in part owing to the high degree
of heterozygosity in diploid apples which, in combination with an ancient genome duplication,
complicated the assembly. Recently, double- and trihaploid individuals have been sequenced,
yielding whole genome sequences of higher quality.[22][23]

The first whole genome assembly was estimated to contain around 57,000 genes,[20] though the
more recent genome sequences support estimates between 42,000 and 44,700 protein-coding
genes.[22][23] The availability of whole genome sequences has provided evidence that the wild
ancestor of the cultivated apple most likely is Malus sieversii. Re-sequencing of multiple
accessions has supported this, while also suggesting extensive introgression from Malus
sylvestris following domestication.[24]

Cultivation

History

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

Wild Malus sieversii apple in Kazakhstan

Central Asia is generally considered the center of origin for apples due to the genetic variability
in specimens there.[25] The wild ancestor of Malus domestica was Malus sieversii, found growing
wild in the mountains of Central Asia in southern Kazakhstan, Kyrgyzstan, Tajikistan,
and northwestern China.[5][26] Cultivation of the species, most likely beginning on the forested
flanks of the Tian Shan mountains, progressed over a long period of time and permitted
secondary introgression of genes from other species into the open-pollinated seeds. Significant
exchange with Malus sylvestris, the crabapple, resulted in populations of apples being more
related to crabapples than to the more morphologically similar progenitor Malus sieversii. In
strains without recent admixture the contribution of the latter predominates. [27][28][29]

The apple is thought to have been domesticated 4,000–10,000 years ago in the Tian
Shan mountains, and then to have travelled along the Silk Road to Europe, with hybridization
and introgression of wild crabapples from Siberia (M. baccata), the Caucasus (M. orientalis),
and Europe (M. sylvestris). Only the M. sieversii trees growing on the western side of the Tian
Shan mountains contributed genetically to the domesticated apple, not the isolated population
on the eastern side.[24]

Chinese soft apples, such as M. asiatica and M. prunifolia, have been cultivated as dessert
apples for more than 2,000 years in China. These are thought to be hybrids
between M. baccata and M. sieversii in Kazakhstan.[24]

Among the traits selected for by human growers are size, fruit acidity, color, firmness, and
soluble sugar. Unusually for domesticated fruits, the wild M. sieversii origin is only slightly
smaller than the modern domesticated apple.[24]

At the Sammardenchia-Cueis site near Udine in Northeastern Italy, seeds from some form of
apples have been found in material carbon dated to between 6570 and 5684 BCE. [30] Genetic
analysis has not yet been successfully used to determine whether such ancient apples were
wild Malus sylvestris or Malus domesticus containing Malus sieversii ancestry. It is hard to
distinguish in the archeological record between foraged wild apples and apple plantations.[31]

There is indirect evidence of apple cultivation in the third millennium BCE in the Middle East.
[31]
There is direct evidence, apple cores, dated to the 10th century BCE from a Judean site
between the Sinai and Negev. [32] There was substantial apple production in European classical
antiquity, and grafting was certainly known then.[31] Grafting is an essential part of modern
domesticated apple production, to be able to propagate the best cultivars; it is unclear when
apple tree grafting was invented.[31]

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

Duration: 1 hour, 1 minute and 36 seconds.1:01:36

Audio 01:01:35 (full text)

The Roman writer Pliny the Elder describes a method of storage for apples from his time in the
1st century. He says they should be placed in a room with good air circulation from a north
facing window on a bed of straw, chaff, or mats with windfalls kept separately. [33] Though
methods like this will extend the availabity of reasonably fresh apples, without refrigeration
their lifespan is limited. Even sturdy winter apple varieties will only keep well until December in
cool climates.[34] For longer storage medieval Europeans strung up cored and peeled apples to
dry, either whole or sliced into rings.[35]

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

Apple cultivars brought as seed from Europe were spread along Native American trade routes,
as well as being cultivated on colonial farms. An 1845 United States apples nursery catalogue
sold 350 of the "best" cultivars, showing the proliferation of new North American cultivars by
the early 19th century.[38] In the 20th century, irrigation projects in Eastern Washington began
and allowed the development of the multibillion-dollar fruit industry, of which the apple is the
leading product.[5]

Until the 20th century, farmers stored apples in frostproof cellars during the winter for their
own use or for sale. Improved transportation of fresh apples by train and road replaced the
necessity for storage.[39][40] Controlled atmosphere facilities are used to keep apples fresh year-
round. Controlled atmosphere facilities use high humidity, low oxygen, and controlled carbon
dioxide levels to maintain fruit freshness. They were first researched at Cambridge University in
the 1920s and first used in the United States in the 1950s.[41]

Breeding

See also: Fruit tree propagation and Malling series

Many apples grow readily from seeds. However, apples must be propagated asexually to obtain
cuttings with the characteristics of the parent. This is because seedling apples are "extreme
heterozygotes". Rather than resembling their parents, seedlings are all different from each
other and from their parents.[42] Triploid cultivars have an additional reproductive barrier in that
three sets of chromosomes cannot be divided evenly during meiosis, yielding unequal
segregation of the chromosomes (aneuploids). Even in the case when a triploid plant can
produce a seed (apples are an example), it occurs infrequently, and seedlings rarely survive. [43]

Because apples are not true breeders when planted as seeds, propagation usually
involves grafting of cuttings. The rootstock used for the bottom of the graft can be selected to
produce trees of a large variety of sizes, as well as changing the winter hardiness, insect and
disease resistance, and soil preference of the resulting tree. Dwarf rootstocks can be used to
produce very small trees (less than 3.0 m or 10 ft high at maturity), which bear fruit many years
earlier in their life cycle than full size trees, and are easier to harvest.[44]

Dwarf rootstocks for apple trees can be traced as far back as 300 BCE, to the area
of Persia and Asia Minor. Alexander the Great sent samples of dwarf apple trees
to Aristotle's Lyceum. Dwarf rootstocks became common by the 15th century and later went
through several cycles of popularity and decline throughout the world.[45] The majority of the
rootstocks used to control size in apples were developed in England in the early 1900s. The East
Malling Research Station conducted extensive research into rootstocks, and their rootstocks are
given an "M" prefix to designate their origin. Rootstocks marked with an "MM" prefix are
Malling-series cultivars later crossed with trees of 'Northern Spy' in Merton, England.[46]

Most new apple cultivars originate as seedlings, which either arise by chance or are bred by
deliberately crossing cultivars with promising characteristics.[47] The words "seedling", "pippin",
and "kernel" in the name of an apple cultivar suggest that it originated as a seedling. Apples can
also form bud sports (mutations on a single branch). Some bud sports turn out to be improved
strains of the parent cultivar. Some differ sufficiently from the parent tree to be considered new
cultivars.[48]
Apples have been acclimatized in Ecuador at very high altitudes, where they can often, with the
needed factors, provide crops twice per year because of constant temperate conditions year-
round.[49]

Pollination

See also: Fruit tree pollination

An apple blossom from an old Ayrshire cultivar

An orchard mason bee on an apple bloom

Apples are self-incompatible; they must cross-pollinate to develop fruit. During the flowering
each season, apple growers often utilize pollinators to carry pollen. Honey bees are most
commonly used. Orchard mason bees are also used as supplemental pollinators in commercial
orchards. Bumblebee queens are sometimes present in orchards, but not usually in sufficient
number to be significant pollinators.[48][50]

Cultivars are sometimes classified by the day of peak bloom in the average 30-day blossom
period, with pollinizers selected from cultivars within a 6-day overlap period. There are four to
seven pollination groups in apples, depending on climate:
  Group A – Early flowering, 1 to 3 May in England ('Gravenstein', 'Red Astrachan')

 Group B – 4 to 7 May ('Idared', 'McIntosh')

 Group C – Mid-season flowering, 8 to 11 May ('Granny Smith', 'Cox's Orange Pippin')

 Group D – Mid/late season flowering, 12 to 15 May ('Golden Delicious', 'Calville blanc

 Group E – Late flowering, 16 to 18 May ('Braeburn', 'Reinette d'Orléans')

 Group F – 19 to 23 May ('Suntan')

 Group H – 24 to 28 May ('Court-Pendu Gris' – also called Court-Pendu plat)

One cultivar can be pollinated by a compatible cultivar from the same group or close (A with A,
or A with B, but not A with C or D).[51]

Maturation and harvest

See also: Fruit picking and Fruit tree pruning

Lauri Kristian Relander, the former President of Finland, with his family picking apples in the
1930s
 

Apples being harvested in Wenatchee, Washington, United States (2010)

Cultivars vary in their yield and the ultimate size of the tree, even when grown on the same
rootstock. Some cultivars, if left unpruned, grow very large—letting them bear more fruit, but
making harvesting more difficult. Depending on tree density (number of trees planted per unit
surface area), mature trees typically bear 40–200 kg (90–440 lb) of apples each year, though
productivity can be close to zero in poor years. Apples are harvested using three-point ladders
that are designed to fit amongst the branches. Trees grafted on dwarfing rootstocks bear about
10–80 kg (20–180 lb) of fruit per year.[48]

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

Crops ripen at different times of the year according to the cultivar. Cultivar that yield their crop
in the summer include 'Sweet Bough' and 'Duchess'; fall producers include 'Blenheim'; winter
producers include 'King', 'Swayzie', and 'Tolman Sweet'.[38]