FRUIT BREEDING: PAST, PRESENT, AND FUTURE1

JULES JANICK
Fruit breeding is an ancient technology with dynamic current technology and an exciting future
(Janick and Moore, 1975, 1996). In its broadest sense, fruit breeding refers to the purposeful genetic
improvement of fruit crops through various techniques including selection, hybridization, mutation
induction, and molecular techniques. Its origins trace to the domestication process in prehistory and
antiquity, where useful species were choses and cultivated, and improved by continuous selection
(Janick, 2005, 2011). Through the millennia genetic improvement of these species have been achieved
by grower selection, first from natural seedling populations and then from grower field that continued
unique genoypes fixed by vegetative propagation. Spontaneous hybridization between wild plants and
cultivated clones was critical to the early domestication of fruits.
Much of the world fruit industry is still based on grower selection from chance seedlings as well
as mutations (sports) and as a result many fruit species are characterized by a narrow germplasm base
(Janick, 2005). These elite seedlings have unique attributes such as outstanding flavor and texture,
high fruitfulness and productivity, but also special problems associated with limited adaptation, and
pest and disease susceptibility. Deficiencies in many cases have been ameliorated through cultural
practices to prop them up including the use of rootstocks, insect and disease control practices, growth
regulators, and special handling and storage technology. While some fruit and nut crops—fig, date, and
almond, for example—are little changed from antiquity. Some fruits, such as peach, have been so
transformed by continuous selection and genetic recombination that they are far removed from their
wild progenitors. Not until the beginning of the 19th century was selection purposely imposed through
cycles of hybridizations and selection. Current progress has been achieved through intensification of
the same forces that have occurred naturally with emphasis on increased adaptability through
hardiness, reducing chill requirements, and photoperiod insensitivity, plus resistance to biotic stress.
(Moore and Janick, 1983). Because many of our fruits are essentially little changed from wild types
continued progress can be expected. Molecular techniques hold out the promise of increasing the

Department of Horticulture & Landscape Architecture, Purdue University, West Lafayette, Indiana, USA
1
This article has appeared in a different version in Temperate Fruit Breeding, Fruit, Vegetable and Cereal
Science and Biotechnology 5 (Special Issue 1), 2011.
efficiency of selection through molecular markers and the direct transfer of useful genes into adapted
genotypes through recombinant DNA (transgene) technology (Folta and Gardiner, 2009).
DOMESTICATION
Fruit tree culture originated occurred in various locations including the Fertile Crescent, Asia, and
the New World in the late Neolithic and Bronze Age. About 8000 years ago, a period known as the
second Neolithic Revolution occurred in the Fertile Crescent that involved a change from villages to
urban communities (Childe, 1958). This development of urban centers is associated with the
development of a settled agriculture. This coincides with the beginning of fruit culture, which involved
a long-term commitment to a unique piece of ground. In the case of the date and olive, a fruit orchard
can remain productive for over a century. It is fruit culture that bonds humans to a particular piece of
land and may be a link associated with the concept of territoriality, the development of city-states, and
eventually nationhood.
Zohary and Spiegel-Roy (1975) proposed that fruit culture, in contrast with mere collection,
originated 4000 to 3000 BCE. Although some information before this period is based on archeological
remains, much of it is by inference and conjecture. Perhaps the earliest pictorial evidence of fruit
growing occurs in a 1 m tall alabaster vessel known as the Uruk vase found in Jemdet Nasr levels at
Uruk that date from about 3000 BCE. Uruk (Erech) is on the Euphrates just north of Basra, Iraq. The
imagery from vase base to rim, depicts water at the bottom, cultivated plants (barley and sesame) and
domestic animals above, followed by attendants bearing baskets of fruit offerings presented to the
Goddess Innana, later known as Istar, in a marriage ceremony to a king (Bahrani, 2002), Unfortunately
the fruits cannot be identified, but they are large and of various shapes. Predynastic drawings in Egypt
depict the date palm.
The development of fruit culture in the Fertile Crescent evolved at two loci: the Tigris-
Euphrates civilization of Mesopotamia and the Nile valley culture of Egypt. Information on the ancient
origins of fruit culture comes from archeological remains of fruit, and from pictorial and literary
evidence. The high culture of Mesopotamia and Egypt produced a rich art in which fruit is a common
motif. A trove of paintings and sculpture, and desiccated remains is found in Egyptian tombs and
monuments. The Sumerian discovery of writing in the 3rd millennium BCE, and Egyptian writings
somewhat later, inaugurated the literary tradition that survives today as a result of the near
indestructibility of the baked clay tablets used for cuneiform script, the wide use of stone carving for
hieroglyphics, and the preservation of papyrus in desert tombs. Later infusions of species and
technology came from Greece, Persia, Turkey, India, China and New World civilizations. By classical
times in Greece and Rome fruit culture had achieved a sophisticated level, not exceeded for over a
millennium.
The first fruits crops to be domesticated appear to be the date palm, olive, grape, almond, fig,
and pomegranate. Asian temperate pome fruits (apple, pear, quince, medlar) and stone fruits
(apricot, cherry, peach, and plum) were fully domesticated by antiquity. Citrus fruits were
domesticated early in China but reached the West in waves starting with citron. Tropical Asian fruits
(mango and banana) and fruits from the Americas (avocado, papapa, and pineapple) were developed in
prehistory. A number of popular fruits and nuts were only domesticated in the 18th to 20th centuries
including various brambles, vacciniums, pecan, and kiwifruit, while many fruits, although extensively
collected and marketed are in the process of domestication (lingonberry, various cacti, durian).
Domestication of fruit crops resulted from selection of elite natural variants with further improvement
arising from recombinants produced by natural intercrosses involving selected types. Cultural
practices often unique to each crop, such as irrigation, pollination control, grafting, (Fig. 1), rootstocks,
pruning and training, storage, and processing, were developed to extend use, increase productivity, and
improve quality.
Fruit Domestication and Genetic Alteration
Fruit crops are characterized by a number of common features. Noteworthy is the obvious
appeal of taste which many consider delicious—often a combination of sweetness, acidity and
aromatic constituents. The desirable taste and colors of many fruits is a naturally selected trait
associated with seed dispersal often mediated by mammal (Steyn, 2012). Most fruit crops are highly
cross-pollinated, and tree fruits generally have long juvenility and long life. Most importantly, some
fruit crops have the ability to propagate vegetatively, by such factors as off-shoots, cuttings, or
nucellar seed. Subsequent progress in the improvement of fruit crops resulted from continual selection
in seedling populations, especially from natural inter-crosses among elite clones or with wild or
introduced clones, that vastly speeded up the process. This process has been efficacious, and in spite of
progress in plant breeding, many grower-selected clones are still being grown.
Domestication of fruits involves a combination of events including species selection, recurrent
selection of elite clones, and vegetative propagation combined with horticultural technology such as
irrigation in dry climates, pruning and training, pollination in the case of date palm, and storage and
processing technology. Genetic changes associated with domestication of fruits (Table 1) include the
breakdown of dioecy, loss of self-incompatability, induction of parthenocarpy and seedlessness,
polyploidy and allopolyploidy, loss of toxic substances, ease of vegetative propagation, and loss of
spines, thorns, or pubescence. Other changes due to selection include increase in fruit size, increase in
sugar content, and increase in storage and shelf life. Factors contributing to genetic improvement
include interspecific recombination, polyploidization, and continued selection involving generations
of sexual recombinants.
Many fruit crops differ from their wild progenitors by a few characters that have appeared as
mutations (Table 2). Typically these mutations are not advantageous to the plant in its natural setting
as they reduce fitness, but would clearly have been immediately selected by humans. The changes from
bitter to sweet seed in almond, and seeded to seedless fruits along with parthenocarpy (banana and
plantain, citrus, fig, grape, persimmon, and pineapple) would have negative fitness but very positive
selective value. Parthenocarpy has two advantages: it eliminates the need for pollination, and is one
path to seedlessness which has proved important in grape, banana, and citrus. In dioecious fruit crops,
mutations inducing hermaphroditism (strawberry, grape, and papaya) are associated with
domestication. Others mutations associated with domestication include loss of spines (brambles,
pineapple, pome fruits, and citrus), loss of fruit pubescence (peach), and changes in growth habit
mutations (pome and stone fruits). In many fruit crops, fruit color mutations (sports) have become
increasingly important, especially in apple, pear, and grapefruit. Some of these mutations are not
heritable because they do not occur in the appropriate meristematic layer.
The development of fruit growing evolved from an interaction of genetic changes and
cultivation technology, often unique for each species (Janick 2005). Some idea of how this has
occurred can best be inferred from the history of two recent domesticates: cranberry and kiwifruit.
What occurred in these crops probably occurred in the past with others, although each crop is unique
with its own set of problems and prospects, and each has its own story. Both cranberry and kiwifruit
were widely appreciated and entered commerce from wild stands long before domestication. The
cranberry had been collected in North America since colonial America, but only became cultivated
in the 19th century. Successful cultivation involved developing a series of practices to grow a plant
adapted to aquatic conditions. The kiwifruit, a dioecious vine native to China has been appreciated
since the 8th century in China and probably much earlier but was never cultivated there. (It was
introduced to England and North America in the beginning of the 20th century, but New Zealand
claims the honor of domestication.) While the plant was introduced to England and the United States,
the plant languished there, emphasizing the key role of champions. A cultivation system worked out
by New Zealand nurserymen and growers involved training and pruning on a trellis, with provision
for pollination. The preferred pistillate and staminate clones (‘Hayward’ and ‘Bruno’, respectively)
were selected from seed introduced into New Zealand from China. After the germplasm was
selected, cultivation techniques established, and markets developed, the technology was quickly
transferred and kiwifruit became a world fruit crop in less than 25 years.
In both cranberry and kiwifruit, the early elite selections of wild plants were of high quality and
could be vegetatively propagated—by cuttings in the case of cranberry and grafting in the case of
kiwifruit. Selection, combined with the ability to fix unique combinations by vegetative propagation,
was the key breeding technique in these two crops, as in all fruit crops. Breeding work has continued
but even after 100 years, the selections made very early still dominate the industry.
In both cranberry and kiwifruit, related species are under consideration as potential new crops.
In kiwifruit, the related yellow-fleshed Actinidia chinensis has been introduced, and the small-fruited,
hardy A. arguta (also known as tara fig) is under consideration as a new domesticate and now widely
planted in northern home gardens. In the vacciniums, two related crops—blueberry (especially
lowbushtypes in Maine) and lingonberry in Sweden—were also widely appreciated and harvested from
the wild, but with remarkably different outcomes. Blueberry had more promise as a commercial fruit
than did cranberry or lingonberry because the fruit could be consumed fresh as well as processed and
there was greater diversity in a number of species. While the domesticates of cranberry and kiwifruit
are little changed from their wild forms, the blueberry has undergone remarkable transformation due
to interspecific hybridization and ploidy manipulation. The culture of blueberry was dependent on the
understanding that the vacciniums are an acid-loving species and required the ammonium form of
nitrogen. Intensive selection and breeding with various species of different ploidy levels transformed
this crop into a relatively large industry of wide adaptation. Lingonberry, on the other hand, a large
Scandinavia export crop from forest-collection, never became domesticated, probably because there
was no shortage of collectable fruit. This crop is still based on merely managed wild plantings.
FRUIT BREEDING
Fruit breeding as an organized activity is a 19th century innovation. Current progress was
achieved through intensification of the same forces that have occurred during domestication with
increasing emphasis on increased adaptability through hardiness, lowered chill requirement,
photoperiod insensitivity, resistance to biotic stress, plant architectural modifications, and selection of
color mutations. Molecular techniques hold out the promise of increasing the efficiency of selection
through molecular markers and insertion of individual genes through transgene technology.
Early beginnings of fruit breeding technology can be demonstrated in strawberry and pear.
The modern strawberry is derived from hybrids between two octoploid (2n=56) native American
species, both usually dioecious: Fragaria virginiana indigenous to the East coast of the North America
but reaching Europe in the 17th century, and F. chiloensis, native from Alaska to Chile. Hybrids
between these two species were produced naturally in Brest, France early in the 18th century when a
pistillate clone of the large-fruited F. chiloensis, introduced by Amédee François Frézier, a French
army officer (and spy) whose family name curiously derives from the French word (fraise) for
strawberry, was inter-planted with staminate plants of F. virginiana. The new hybrids (now known as
F. ×ananassa or pineapple strawberry, after their shape and aromatic flavor, initiated the modern
strawberry industry. Selection through the years has resulted in tremendous changes as the plant
evolved from a predominantly dioecious species with male and female plants into a hermaphroditic
species, in which flowers contained both stamens and pistils. The development of the day neutral
character from interspecific crosses has resulted in continuous fruit production and has transformed the
industry. Modern breeding has greatly increased fruit size and firmness.
 Systematic breeding of European pear was first carried out by Jean Baptiste Van Mons
(1765–1842), a Belgian physician, pharmacist, and physicist and an early apostle of selection in plants.
He collected clones of pear and sequentially planted (open pollinated) seed of the best material making
new selections for eight generations. An early fruit book, The American Fruit Culturist (1863) by John
J. Thomas (1810–1895) states that the mean time from seed planting to fruiting in the first cycle was
12 to 15 years, 10 to 12 in the second cycle, 8 to 10 by the third, 6 to 8 by the fourth, and 5 by the
fifth. By the 8th generation several fruit trees fruited at the age of four years (emphasis by the author,
presumably based on correspondence of Von Mons). This may be the first example of data on long-
term selection in plants. Recent breeding efforts have concentrated on fireblight resistance.
Thomas Andrew Knight (1759–1838) was the first to improve fruits by selection from
genetic recombination derived from inter-pollinations of clones (Fig. 2). An early proponent of the
development of plant improvement through crossbreeding and selection he literally initiated the field of
fruit breeding. He released a number of improved fruit cultivars (apple and pear, cherry strawberry, red
currant and strawberry, and cherry, nectarine, and plum). His studies on the effects of pollen in the
garden pea on seed characters presaged the work of Gregor Mendel carried out 40 years later. He
describes dominance and segregation, although he failed to make the brilliant leap of Mendel in
relating phenotypic characters to the factors we now know as genes. Gregor Mendel, the father of
genetics, was also involved in designing apple and pear breeding programs.
In the United States, fruit breeding became a part of research at the state and federal
experiment stations and many breeding programs were initiated throughout the United States.
Practically each state had fruit breeding programs at one time. Fruit breeding also became an activity of
the private sector. An American nurseryman, Luther Burbank (1849–1926), was the first to consider
fruit breeding as a commercial endeavor, and although he distrusted Mendelism, he was a staunch
believer in the evolutional theories of Darwin. Burbank was also influential in obtaining passage of
the Townsend-Purnell Plant Patent Act signed in 1930 which initiated the protection of fruit cultivars.
At present private breeders are an important part of breeding efforts in Prunus (especially peach and
plum), strawberry, and raspberry.
 Although organized programs of fruit breeding by universities, state and federal experiment
stations have been a major activity since early in the 20th century, the results have been variable and
vary from ineffectual to extraordinarily successful. The peach is an example where breeding efforts
have had great impact. For example breeding peaches with low chill requirements as well as increased
hardiness has greatly expanded the areas where this crop can be successfully produced. The
development of firm fleshed peaches became the basis for a large processing industry in California.
Much of the world fruit industries are still based on grower-selected clones. The reason for the
lack of progress of fruit breeding programs is twofold. First, vegetative propagation permits the genetic
fixation of naturally occurring variation. Because of the vast populations involved in seedling orchards,
the quality of the selected clones over hundreds and even thousands of years of selection is very
high. Second, the difficulties and expense inherent in fruit breeding have inhibited long term breeding
efforts. Progress from breeding a number of fruit crops, however, has shown significant advances in
the second half of the 20th century and selections from controlled crosses are increasingly important
in many crops. In apple, although chance seedlings such as ‘Delicious’ and ‘Golden Delicious’ have
long dominated the world market. ‘Fuji’, a seedling derived from a Japanese breeding program (‘Ralls
Janet’ × ‘Delicious’) is now the leading world cultivar.
Hybridization and Selection
Selection from sexual recombinants is still the dominant force in the domestication process as well as
modern fruit breeding. The isolation of elite selections, combined with mass plantings created a
situation where mass selection and recurrent selection could operate naturally. This has recently been
confirmed in apple, where elite selections from Kazakhstan are very close to cultivated varieties
(Harris et al. 2002).
Selection from sexual recombination in apple can be clearly followed in North America,
now considered a secondary center of origin (Janick et al., 1996). In colonial America, starting in the
1600s, the apple was imported by immigrants, some as scions but most as seeds from Holland,
Germany, France, and the British Isles (Beach 1905). Pioneers were encouraged to plant apples and the
requirement for settling Ohio (1787–1788) included that the settler must harvest at least 50 apple or
pear trees and 20 peach trees (Morgan and Richards 1993). Apples, once introduced, were carried far
into the wilderness by Native Americans, traders, and missionaries and became naturalized. In 1806,
Jonathan Chapman (the legendary Johnny Appleseed), distributed apple seeds from cider mills in
western Pennsylvania and founded a nursery in West Virginia. The apple flourished in the new
territories with the greatest use for hard cider, the distilled product called apple jack, and vinegar for
preservation of fruits and vegetables. Many of the imported apple clones were unadapted, and the
selection of natural seedlings from orchards became the glory of 19th century American pomology. In
1905, 698 apple cultivars were described in Beach’s (1905) Apples of New York. In the United States
the screening of open-pollinated, chance seedlings resulted in thousands of selections, many of which
proved to be outstanding cultivars, including ‘Golden Delicious’, ‘Delicious’, ‘Jonathan’, ‘McIntosh’,
‘Rome Beauty’, ‘York Imperial’, ‘Stayman Winesap’, ‘Yellow Newtown’, ‘Winesap’, ‘Rhode Island
Greening’, ‘Northern Spy’, and ‘Gravenstein’. These unique selections selected from thousands of
growers became the starting point for hybridization programs in the early history of apple breeding.
th
‘Golden Delicious’ had a profound influence on apple growing in Europe in the 20 century and
further proved to be a prepotent parent producing many important new cultivars from breeding efforts.
Differences in tolerance to many pests are observed in any large collection of apples.
Immunity to apple scab has been transferred from Malus floribunda by backcrossing, but the
problem of races has also appeared in some areas. This places the durability of this gene (Vf) in
question, although it has held up for many years in the United States (Janick 2002).
Conventional plant breeding based on continuous selection of superior phenotypes from
genetically variable populations derived from sexual recombination is powerful because it is
evolutionary. Progress can be cumulative with improved individuals continually serving as parents for
subsequent cycles of breeding. Thus, genetic improvement has made substantial changes in a number
of fruit species when the effort has been well supported and long term.
The objectives of fruit breeding varied with location. In northern locations objectives were
based on hardiness. Breeding for apple hardiness is an essential character in the northern US and in
Scandinavia. In southern locations low chill requirements received attention. Breeding for low chill has
greatly increased the range of peach and nectarine in subtropical areas. Throughout the humid parts of
the country disease and pest resistance were important objectives. In the western US, the main
characters being selected were firmness and amenability to long distance shipping. Selection for fruit
quality, based on flavor, color, and shelf life, and texture was the goal of modern fruit breeding
programs but this factor often was sacrificed for appearance and shipping quality. Improved fruit
texture has been important in apple and peach. Where processing was important to the industry,
processing quality became key. Many processing industries soon came to be dependent on unique
cultivars with special qualities such as ‘Bartlett’ pear. The processing industry of pineapple was long
based on a single cultivar. ‘Cayenne’ and its spineless sport (‘Smooth Cayenne’) that was uniquely
adapted to producing canned slices. However, organoleptic quality is increasingly being considered as
key if fruits for the fresh market. A sweeter, yellow-fleshed seedling of pineapple resulting from a
hybridization in Hawaii. PRI 73-114 (‘MD-2’) is now being marketed as “Del Monte Gold” and has
transforming the world fresh fruit industry because of better fresh fruit quality and appearance than
‘Smooth Cayenne’.
BIOTECHNOLOGY AND FRUIT BREEDING
There are a number of restraints to conventional fruit breeding which are especially limiting
in tree fruits with their long juvenile period, large plant size, and which are represented by unique,
highly-selected heterozygous genotypes. These restraints include: (1) the reliance on naturally
occurring variation which may be unavailable, or more likely found in very primitive and unadapted
genotypes; (2) restraints due to the inability of the sexual system to incorporate genes from non-
related species and especially the inability of the sexual system to incorporate small changes without
recombination resulting in the loss of unique combinations; (3) problems with incompatibility and
sterility; (4) the randomness of induced mutation; (5) the difficulty of selection in detecting infrequent
or rare recombinants, (6) and the dependence of conventional breeding on time to generate cycles of
recombination; and space to grow the necessary populations to recover superior recombinants; and
resources to be able to select, identify , and evaluate desirable recombinants.
The biotechnological revolution in the last half of the 20th century is based upon novel
genetic strategies derived from microbial and molecular techniques including, in vitro propagation,
embryo rescue, protoplast fusion, marker-assisted selection and recombinant DNA (transgene)
technology. Advances in molecular genetics may overcome some of the limitations to conventional
fruit breeding by increasing selection efficiency using molecular markers and by transgene technology
whereby individual genes from various sources may be inserted without disturbing unique genetic
combinations. Biotechnology has reinvigorated the art and science of fruit breeding. For example,
embryo rescue techniques have been a key factor in the breeding of seedless grapes. However, the
great hopes for marker-assisted selection and transgene technology has yet to reach its potential.
Marker-assisted Selection
Advances in genomics, the study of the DNA structure of genomes, has led to complete genetic
mapping of many crop plants including fruits species. Efforts are underway to map the genomes of
many fruit species with special emphasis on the Rosaceae (Folta and Gardiner 2009). Full genomic
sequencing in peach, apple, and strawberry has made it possible to locate useful genes include those
affecting quantitative traits. The combination of advances in informatics, sequencing technology, and
genomics makes it theoretically possible to select on the basis of the genotype instead of the
phenotype. This has potential benefits to fruit breeding where the cost of phenotype selection are
very high because of the long time necessary for fruiting and the large amounts of land required for
each seedling.
Transgene Technology
This innovation theoretically offers many advances for fruit breeding because it allows discrete
changes to be made of well-adapted genotypes. For example it would be possible to induce such
characters as fireblight resistance into ‘Bartlett’ pear, uniquely suited for processing, without changing
any of the quality characters. Progress has already been achieved in incorporating virus resistance in
papaya (Gonsalves et al. 2006). However, fears of consumer resistance has been a problem when
introduced genes are considered from widely divergent species. However, syteny in Rosaceae (Arús et
al. 2006) suggest that the transfer of genes within the family will partially calm the fears of
consumers. However, at the present time it is clear that the future of fruit breeding lies in the use of
these new techniques as a tool to enhance but not replace traditional methods.
Patents and the Privatization of Fruit Breeding
In the 20th century most organized fruit breeding programs were carried out with free release of
germplasm. The advantages of this practice were cooperation among fruit breeders and especially the
sharing of germplasm. However, because fruit breeding turned out to be an expensive activity most
experiment stations slowly receded from this activity as the competition among institutions began to
be spurred by the funds generated by granting institutions favoring more basic research. In the United
States this led to severe decrease in breeding activity. However, the increase in patent protection
indicated that fruit breeding could be made self supporting. This was made evident by the large
royalties generated by the strawberry breeding program of the University of California. The rise in
patent protection has also invigorated a number of private breeding programs.
The situation was made more complex when the concept of club cultivars was developed (Janick
,1994). Under this system, with the protection of patents, unique cultivars could be released to a small
selection of growers so that quality could be controlled and supply could be limited. Under this system
the profits from plant breeding extended to the revenues generated by the fruit crop each year where
heretofore it was generated merely by the sale of plants. Under this system, revenue of superior
genotypes could be very much extended to the life of the plant itself.
THE FUTURE OF FRUIT BREEDING
Fruit breeding has been in a state of flux because the traditional support by government
experiment stations received throughout the world has declined as a reaction to the high costs of long
term programs that are necessary. However, the globalization of the fruit industry has underscored
the economic value of improved germplasm indicating that support for fruit breeding needs to be
increased. Returns from patents offer one solution to impede the decline in funding and to encourage
breeding effort. Patents are essentially a direct tax on the industry to improve breeding and also offer
incentives to breeders when they share in patent royalties. In some countries the costs of foreign patents
has encouraged investment in local breeding programs.
The objectives of breeding efforts in the future must be twofold. The most important is to
increase quality in order to increase consumption. This is important to reverse the decline in per capita
fruit consumption in developed countries. The second effort must be to increase the efficiency with
special emphasis on productivity, annual bearing, and pest resistance. However, the future of fruit
breeding is crop specific. For example the tremendous advances in seedless table grapes have
intensified breeding efforts. In contrast, the breeding of wine grapes is impeded by the association of
traditional cultivar name with the name of the wine. As a result the wine industry has not encouraged
cultivar change except for clonal selection. Practically no advances have been made in the genetic
improvement of banana but there are major projects via international efforts (International Network for
the Improvement of Banana and Plantain or INIBAP). The lack of diversity in export banana makes
breeding essential due to disease pressure (such as sigatoka) but seedless banana is notorious difficult
to hybridize.
Similarly, citrus industry has increased in efforts to produce new seedless, easy-peel clones
while the threat of diseases such as hanglongbing (citrus greening) is an upcoming problem that may
require a genetic solution. The success of breeding in Prunus has encouraged a number of private
breeding organizations especially vibrant in peach and plums. Recent studies of a partially stoneless
plum described by Luther Burbank suggest that the hard stone might be eliminated in Prunus (Callahan
et al. 2009). Similarly the success of breeding efforts to produce late ripening cherries in British
Columbia, Canada has indicated the high economic returns made possible by sustained breeding
effort. Apple breeding has produced many new cultivars and the success of ‘Fuji’ and ‘Honeycrisp’
has underscored the returns from improved quality. There are large efforts in breeding loquat in China
including triploid, seedless clones (Lin et al. 2007). Recent success with blackberry with the
introduction of thornlessness, primocane fruiting, reduced seediness, and large, tasteful fruit has created
a new global industry (Finn and Clark, 2011).
In the case of underutilized or new fruit corps, knowledge of domestication should be used to
predict future changes. Thus one might anticipate that in kiwifruit, hermaphroditic mutations would
eliminate the need for staminate clones as pollinators, and that fruit skin mutations or breeding could
lead to non-pubescent clones, with more attractive and more edible fruit surface. The use of
interspecific hybridization should lead to improvement Prunus and Rubus. A number of tropical fruits
are candidates for commercialization, provided postharvest technology can be improved. One of the
likely candidates for domestication is pitaya (species of Selenicereus, Hylocereus, and Cereus), an
extremely attractive fruit of columnar cactus, but breeding efforts to improve quality is required, since
many selections have been insipid (Mizrahi et al., 2002). In the future, the greatest progress in fruit
breeding will be the improvement of underutilized fruits such as jaboticaba in Brazil.
 Most fruit crops are not far removed from wild species, some perhaps by only a few
generations, so that in most cases continued progress should be possible. Yet, many of our fruit
cultivars are ancient and based on grower-selected seedlings and somatic mutations. In conventional
breeding systems, elite clones are selected followed by fixation by vegetative propagation, and these
provide the raw material for subsequent cycles of hybridization and selection. The development of fruit
culture is based on an interaction between improved clones and cultural practices. Deficiencies are
compensated by such techniques as artificial pollination, the use of disease-resistant and size
controlling rootstocks, extensive methods of disease control, involving complex schedules of pesticide
application, the control of fruit size and annual bearing by manual and chemical fruit and flower
thinning, the control of fruit abscission with growth regulators, and extensive pruning and training
systems. The dilemma is to determine if breeding emphasis should concentrate on quality for
consumers or rather to overcome production problems for growers by genetic solution. Obviously both
must be objectives in modern breeding programs. Because fruit breeding depends on genetic diversity
the development of germplasm resources must be strengthened by collections in situ. In some cases
these collections suggest that a number of fruits that have been ignored, such as quince, are candidates
for increased breeding efforts.
Finally advances in biotechnology must be attached to breeding efforts. Marker assisted
selection could offer significant help in disease and pest resistance breeding but much genetic work
must still be pursed to locate these quality traits. The U.S. Department of Agriculture funded program
founded in 2009 called RosBREED is underway to translated progress in marker-assisted breeding in
rosaceous fruit crops including apple, peach, sweet and tart cherries and strawberry). Similarly
transgene technology could find a place in fruit breeding and the example of virus resistant papaya has
demonstrated that success can be achieved. However, the rationale for transgene technology assumes
that the present germplasm is optimum. This is a conservative approach somewhat similar to backcross
breeding but it is true that many high quality fruits are difficult to replace.
LITERATURE CITED
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Table 1 - Origins and changes associated with domestication and breeding of popular fruits (Janick, 2005).
Species Chromosome Changes associated with
Fruit crop no. Family Origin Flowering in wild species domestication and breeding
Almond Prunus dulcis x=8, Rosaceae SW Asia Hermaphroditic, self- “Sweet” seed, increased kernel
2n=16, 32 incompatible size, self-fertility
Apple Malus × domestica Rosaceae Central Asia Hermaphroditic, self- Combination of size aroma, loss of
x=17, 2n=34, 51 incompatible astringency, sweetness,
parthenocarpy, triploidy
Apricot Prunus armeniaca x=8, Rosaceae Central & E. Asia Hermaphroditic, self- Increase fruit size
2n=16 incompatible
Cherry-sweet Prunus avium x=8, Rosaceae Central Europe & Hermaphroditic, self- Self-compatibility
2n=16 W. Asia incompatible
-tart P. cerasus x=8, 2n=16,
32
Fig Ficus carica x=13, Moraceae E. Mediterranean Gynodioecious Parthenocarpy
2n=26 basin
Loquat Eriobotrya japonica Rosaceae China Hermpahroditic, some Increased size, seedless triploids
x=17, 2n=34, 51 incompatability
Olive Olea europea x=23, Oleaceae Mediterranean Andromonecious Increased fruit size, high oil
2n=46 basin
Peach Prunus persica x=8, Rosaceae China Hermaphroditic Freestone, low chill, fuzzless
2n=16 (nectarine), increased size
Pear Pyrus communis, Rosaceae Central and East Hermaphroditic, self- Combination of size, aroma, loss of
P. pyrifolis, Asia incompatible astringency, sweetness,
P. bretscheiderii, P. parthenocarpy, triploidy
 ussuriensis x=17,
2n=34, 51
Plum Prunus domestica x=8, Rosaceae Europe Hermaphroditic, self- Hexaploid after interspecific
2n=48 incompatible hybridization
Prunus salicina x=8, China
2n=16, 32
Prunus Americana x=8, North America
2n=16
Subtropical and
tropical fruits
Avocado Persea americana Lauraceae Tropical America Hermaphroditic, High oil, smaller seed size
x=12, 2n=24 synchronous protogynous
dicogamy
Banana, plantain Musa sapientum Musaceae SE Asia Monoecious Seedlessness, parthenocarpy,
triploidy
Citrus (orange, Citrus spp. x=9, 2n=18 Rutaceae Southeast Asia, Hermaphroditic Nucellar embryony, interspecific
mandarin, lemon, China hybridization, parthenocarpy
lime, pumello,
grapefruit)
Date palm Phoenix dactilifera Arecaceae S. Mediterranean Dioecious Offshoot production, increased fruit
x=18, 2n=36 basin size
Mango Mangifera indica x=20, Anacardiaceae E. Asia Hermaphroditic Nucellar embryony, loss of fibers in
2n=40 fruit
Persimmon Diospyros kaki x=15, Ebenaceae China Polygamodioecious Loss of astringency, parthenocarpy
2n=90
Papaya Carica papaya x=9, Euphorbiacea Tropical America Dioecious Polygamodioecious, reduced fruit
2n=18 size
Pineapple Ananas comosus x=25, Bromeliaceae Tropical America Hermaphroditic Parthenocarpy, seedlessness
2n=50
Berry and Vine
fruits
Blackberry Rubus spp. x=7, Rosaceae N. America Hermaphroditic Interspecific hybridization,
2n=28,35,42,56,84 polyploidy, thornlessness
Blueberry Vaccinium spp. Ericaceae N. America Hermaphroditic Increased fruit size, interspecific
hybridization, polyploidy
Cranberry Vaccinium Ericaceae E. United States Hermaphroditic Unchanged
macroacarpon
Grape Vitis vinifera x=19, Vitaceae W. Asia Dioecious Hermaphroditic, increased berry
2n=38 size, parthenocarpy
Kiwifruit Actinidia delicious Actinidiaceae China Dioecious Increased fruit size
x=29, 2n=174
A. sinensis x=29, 2n=58
Lingonberry Vaccinium vitis-idaea Ericaceae Circumboreal Hermaphroditic Unchanged
Raspberry Rubus ideaus, Rosaceae Europe, America Hermaphrodictic Interspecific hybridization,
R. occidentalis x=7, polyploidy
2n=14
Strawberry Fragaria ×ananassa Rosaceae Americas Dioecism Hermaphroditic, interspecific
x=7, 2n=56 hybridization
Other spp. 2n=14, 28
Table 2. Genetic changes associated with domestication in fruit crops.
Breakdown of dioecy: fig, grape, papaya, strawberry (unchanged, date palm, kiwifruit)
Loss of self-incompatibility: cherry
Parthenocarpy and seedlessness: apple, banana, blackberry, citrus, fig, grape, pear, persimmon, pineapple,
plantain
Allopolyploidy: banana, plantain, blackberry, raspberry, blueberry, citrus, tart cherry,
European plum, strawberry
Polyploidy
Triploidy apple, banana and plantain, pear
Tetraploidy tart cherry, raspberry, blackberry, blueberry, kiwifruit (Actinidia sinensis)
Hexaploid European plums, kiwifruit (A. deliciosa)
Octaploid strawberry
Loss of toxic substances
“Sweet” seed: almond
Nonastrigency: apple, pear, persimmon, pomegranate
Ease of vegetative propagation
Offshoots: date palm
Rooting: apple (rootstock)
Nucellar embryony: citrus, mango
Loss of spines, thorns, or pubescence: apple, brambles, citrus, peach, pear, pineapple
Figure 1 - Grafting portrayed in a Roman mosaic, 3rd century CE. Source: St. Roman-en-gal, Vienne, France.
Figure 2 - Thomas Andrew Knight (1759–1838).