1. Fruit Breeding: Past, Present, and Future
Jules Janick
Department of Horticulture
Purdue University
West Lafayette, Indiana

2. Time Frame: Agriculture & Fruit Culture
10,000
Neolithic Revolution
Discovery of Agriculture
8000–5000
Second Neolithic Revolution
Villages to Urban communities
Origin of fruit culture
5000–3000
Bronze Age
Horticultural arts (irrigation, vegetative propagation, processing, selection)
3000–2000
Greek and Roman Antiquity
Sophisticated fruit culture

3. Gathering Grain, 4000 BCE. Tassili n’Ajjer, Algeria.
Date palm flanked by gazelle, in Egypt, 4000–3000 BCE

4. The Uruk Vase, Late 4th Millennium BCE
This is the Uruk Vase, made over 5000 years ago, from the biblical city of Erech (near Basra) in present day Iraq. It was one of the treasures missing from the Bagdad Museum that has recently been restored. In a wedding feast an attendant offers a basket of fruit to the Goddess Inana (later known as Ishtar). Horticulture becomes part of religious practice.

5. Grain & Fruit Production in Egypt, 1279 BCE

6. Our fruits are gifts of diversity, not results of systematic breeding, but to unique selection events by unsung and unremembered farmers.
Vincenzo Campi 1580

7. Origin of Fruit Crops Mediterranean basin
Date palm, fig, grape, olive, pomegranate
Europe
Plum, raspberry
Central Asia
Pome fruits, apricot
East Asia
Banana, citrus, peach, kiwifruit, mango, persimmon
Africa
Sycomore fig, coffee
Americas
Avocado, blackberry, papaya, pineapple, strawberry, Vacciniums

8. Fruit Domestication Recognition of species
Selection of elite genotypes
Vegetative propagation technology
Cultivation technology
Pruning and training, irrigation, pollination, pest control
Utilization
Storage, drying, fermentation, processing
Dispersal of fruit crops with human migration
Most fruit crops are closer to wild species than annual crops such as grains

9. Elite selections from Kazakhstan
Commercial Apples
Elite selections from Kazakhstan

10. 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 & seedlessness
Apple & pear, banana & plantain, citrus, fig, grape, loquat, persimmon, pineapple

11. Allopolyploidy
Banana & plantain, blackberry & raspberry, blueberry, tart cherry, European plum, strawberry
Triploidy: banana and plantain, apple, pear
Tetraploid : tart cherry, raspberry, blackberry, blueberry, kiwifruit (Actinidia sinensis)
Hexaploid: European plums, kiwifruit (A. deliciosa)
Octaploid: strawberry

12. Loss of toxic substances
“Sweet” seed: almond
Non-astrigency: 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

13. Recent Domesticates Kiwifruit Vacciniums Cranberry Blueberry
Lingonberry

14. Kiwifruit

15. Cranberry and Lingonberry

16. Blueberry

17. Genetic Structure of Fruit Crops
Cross pollination
High heterogeneity
Asexually propagated
Large size
Long juvenility
Polyploid
Self-incompatible
Sterility and seedlessness
Apomictic

18. Origin of Fruit Breeding
Jean Baptiste Van Mons (1765–1842)
Thomas Andrew Knight (1759–1838)

19. Origin of Fruit Breeding
Gregor Mendel
(1822–1884)
Luther Burbank (1849–1926)

20. Levels of Fruit Breeding
Primitive
Vegetative propagation of unique clones
Mass culture of superior clones
Selection of chance seedlings
Propagation of clonal variants
Conventional Breeding
Recombination of elite clones
Interspecific recombination
Backcross breeding
Mutation breeding
Biotechnology
Embryo rescue
Paraplast fusion
Marker assisted selection
Transgenic breeding

21. Effects of Organized Fruit Breeding on the Commercial World Industry
Negligible
Slight
Moderate
Major
Banana & plantain
Chestnut
Date palm
Fig
Grape (wine)
Lingonberry
Olive
Pomegranate
Tart cherry
Citrus
Cranberry
Hazelnut
Kiwifruit
Papaya
Persimmon
Pear European
Pineapple
Almond
Apple
Apricot
Avocado
Pear Asian
Pecan
Blueberry
Brambles (raspberry & blackberry)
Cherry (sweet)
Currants
Grape (table)
Strawberry
Peach & nectarine
Plum

22. Advantages of Conventional Fruit Breeding
Evolutionary
Progress is cumulative—improved selections serve as parents
Transgressive
Offspring can exceed either parent

23. Limitations of Conventional Breeding
Technical
Expensive
Long juvenile period
Large plants expensive to maintain
Difficulty of selection
Inability to preserve unique genotypes
Genetic recombination uncontrollable
(no way to induce small changes)
Linkage of desirable and undesirable traits
Restricted to natural-occurring variation or
random mutations

24. Limitations of Conventional Breeding
Nontechnical
Must compete with seedlings found
world-wide
Market resistance to new cultivars
Patent costs and restrictions
Testing problems

25. Advantages of Transgene Technology
Allows insertion of single genes without disruption of genotype
Not limited by sexual barriers
Multigene transfer possible
Permits selection by molecular markers (independent of development or emvironment)
Bioengineering (vaccines, plastics, oils)

26. Limitations of Transgene Technology
Technical
Transformation impediments
Selection of transgene
Identification and isolation of transgenes
Chimeral problems
Expression
Shortage of suitable promoters
Testing problems

27. Limitations of Transgene Technology
Nontechnical
Legal problems
Consumer resistance

28. Banana

29. Evolution of Cultivated Bananas

30. Pineapple

31. Propagation of Pineapple

32. Spineless Mutation

33. Del Monte Gold

34. Citrus
Mandarin

35. Nucellar Embryony

36. Color Sports in Grapefruit

37. Interspecific Hybridization in Citrus
(Orange)
(Mandarin)
(Grapefruit)

38. Tangerine × Grapefruit
Tangelo
Minneola ×
Page

39. Tangerine × Orange Tangor
Murcott

40. Strategies for Breeding Seedless Citrus
Ploidy Manipulation (French system)
Exploit Nonreduction in diploids Chomosome counts or flow cytometry Exploit Small seeds with embryo rescue
Diploid × Tetraploid crosses
Protoplast Fusion Produce haploids by irradiated pollen x+2x somatic hybridization and embryo regeneration
Regeneration of triploid endosperm
Induce Sterility by Irradiation (Israeli system)

41. Achievements at CIRAD/INRA of triploid breeding of citrus with the 2x × 4x strategy

42. Stone Fruits (Prunus)
Peach
Apricot
Almond
Plum
Cherry

43. Peach from Pompeii

44. Stoneless Plum (Callahan Dardick Scorza JASH 1934 2009)

45. Synteny in Prunus & Malus

46. Pome Fruits (Apple, Pear)

49. Barrel Sprayer

50. History of the PRI Program

51. Dan Dayton
Jules Janick
Ed Williams
Fred Hough
Ralph Shay

52. Sources of Scab-resistance

53. Sources of Resistance Dominant Resistance Multigenic Resistance
M. atrosanguinea (2 genes)
M. baccata jackii
Dolgo
M. floribunda 821
Geneva
Jonsib
M. micromalus
M. prunifolia
M. prunifolia microcarpa
M. prunifolia xanthocarpa
Morton Arboretum 4, 8, 16, 1255
Russian A (2+ genes)
Multigenic Resistance
Antonovka
M. baccata (selected seedlings)
M. sargentii 843
M. sieboldii
M. toringo 852
M. zumi calocarpa

54. Defined Gene Pools Vf M. floribunda Vm M. micromalus (pit) Vr Russian
Vbj M. baccata jackii
Vb Hansens’s baccata #2
Va Antonovka (pit)

57. Transferring Vf by Backcrossing

58. PRI Releases Prima 1970 Priscilla 1972 Sir Prize 1975 Jonafree 1979
Redfree 1981
Dayton 1988
Williams’ Pride 1988
Enterprise 1993
Pristine 1993
GoldRush 1993
Scarlett O’Hara 2000
Pixie Crunch 2002
Sundance 2002
CrimsonCrisp 2005
Joint Releases
Viking (Wisc.) 1969
Priam (France) 1974
McShay (Ore.) 1988
Primiera (Italy) 1995
Nambu (Japan) 1994
Constance (Ger.) 1995
Primivera (Can.) 1996
Juliet (France) 1999

59. Prima
Redfree
Williams’ Pride
Sundance
Pristine
Enterprise
GoldRush
Pixie Crunch
CrimsonCrisp

60. Races of Venturia inaequalis
Race Source Susceptible Material
1 Worldwide Most of the world’s cultivars
2 South Dakota, USA M. baccata, ‘Dolgo’, ‘Alexis’, ‘Bittercrab’ segregates of R A, ‘Geneva’
3 Nova Scotia, Canada ‘Geneva’
4 Lafayette, IN, USA Segregates of R A
5 Norwich, England Micromalus pit type resistance, M. atrosangunia 804
6 Ahrensburg, Germany ‘Prima’ (Vf cultivars) but not Evereste’ M. × ‘Perpetu’ and M. floribunda 821
7 England and Europe M. floribunda 821

61. Loquat (Eriobotrya japonica) Subtropical Pome Fruit

62. Triploid Loquat Seedling ploidy No. seedlings % Diploid 44,828 99.31
225
0.50
Tetraploid 50
0.11
Pentaploid 10
0.02
Mixoploid 26
0.06

63. Seeded diploid loquat as compared to seedless triploid (courtesy of G
Seeded diploid loquat as compared to seedless triploid (courtesy of G. Liang)

64. Molecular linkage map obtained from ‘Algerie’ and ‘Zaozhong-6’

65. Papaya Resistant to PRSV (D. Gonsalves)
Transgenic ‘SunUp’-Hawaii (CP/CP)
Transgenic ‘Rainbow’-Hawaii (CP/-)
Transgenic ‘Khaknuan’ Thailand
Tolerant ‘Thapra 2’ Thailand

66. Conclusions
Our fruits are legacies of Neolithic and Bronze Age farmers
Persistence of farmer-selected cultivars due to unique quality factors making them difficult to replace
Deficiencies made up by cultural techniques and genetic changes

67. Future Advances:
Search for specific mutations (parthenocarpy and seedlessness, breakdown of dioecy, loss of compatibility, high sugar, loss of astringency)
Use of interspecific crosses to create new fruits (Citrus, Prunus, Rubus)
Selection of underexploited germplasm (pitaya)
Induction of seedlessness
Transfer of genetic resistance to diseases and pests
Increased use of biotechnology (marker assisted selection, new traits, early flowering)
Emphasis on consumer satisfaction