Fruit Breeding: Past, Present, and Future Jules Janick Department of Horticulture Purdue University West Lafayette, Indiana 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) Sophisticated fruit culture 3000–2000 Greek and Roman Antiquity Gathering Grain, 4000 BCE. Tassili n’Ajjer, Algeria. Date palm flanked by gazelle, in Egypt, 4000–3000 BCE The Uruk Vase, Late 4th Millennium BCE Grain & Fruit Production in Egypt, 1279 BCE Our fruits are gifts of diversity, not results of systematic breeding, but to unique selection events by unsung and unremembered farmers. Vincenzo Campi 1580 Origin of Fruit Crops Mediterranean basin Date palm, fig, grape, olive, pomegranate Europe Central Asia East Asia Plum, raspberry Pome fruits, apricot Banana, citrus, peach, kiwifruit, mango, persimmon Sycomore fig, coffee Avocado, blackberry, papaya, pineapple, strawberry, Vacciniums Africa Americas 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 Commercial Apples Elite selections from Kazakhstan Genetic Changes Associated with Domestication in Fruit Crops Breakdown of dioecy Fig, grape, papaya, strawberry (unchanged, date palm, kiwifruit) Loss of selfincompatibility Cherry Parthenocarpy & Apple & pear, banana & plantain, seedlessness citrus, fig, grape, loquat, persimmon, pineapple 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 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 Recent Domesticates Kiwifruit Vacciniums Cranberry Blueberry Lingonberry Kiwifruit Cranberry and Lingonberry Blueberry Genetic Structure of Fruit Crops Cross pollination High heterogeneity Asexually propagated Large size Long juvenility Polyploid Self-incompatible Sterility and seedlessness Apomictic Origin of Fruit Breeding Jean Baptiste Van Mons (1765–1842) Thomas Andrew Knight (1759–1838) Origin of Fruit Breeding Gregor Mendel (1822–1884) Luther Burbank (1849–1926) Levels of Fruit Breeding Primitive Vegetative propagation of unique clones Mass culture of superior clones Selection of chance seedlings Propagation of clonal variants Conventional Recombination of elite clones Breeding Interspecific recombination Backcross breeding Mutation breeding Biotechnology Embryo rescue Paraplast fusion Marker assisted selection Transgenic breeding 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 Advantages of Conventional Fruit Breeding Evolutionary Progress is cumulative—improved selections serve as parents Transgressive Offspring can exceed either parent 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 Limitations of Conventional Breeding Nontechnical Must compete with seedlings found world-wide Market resistance to new cultivars Patent costs and restrictions Testing problems 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) Limitations of Transgene Technology Technical Transformation impediments Selection of transgene Identification and isolation of transgenes Chimeral problems Expression Shortage of suitable promoters Testing problems Limitations of Transgene Technology Nontechnical Legal problems Consumer resistance Banana Evolution of Cultivated Bananas Pineapple Propagation of Pineapple Spineless Mutation Del Monte Gold Citrus Mandarin Nucellar Embryony Color Sports in Grapefruit Interspecific Hybridization in Citrus (Orange) (Mandarin) (Grapefruit) Tangerine × Grapefruit × Tangelo Minneola Page Tangerine × Orange Tangor × Murcott Strategies for Breeding Seedless Citrus Ploidy Manipulation (French system) 1. Exploit Nonreduction in diploids Chomosome counts or flow cytometry Exploit Small seeds with embryo rescue 2. Diploid × Tetraploid crosses 3. Protoplast Fusion Produce haploids by irradiated pollen x+2x somatic hybridization and embryo regeneration 4. Regeneration of triploid endosperm Induce Sterility by Irradiation (Israeli system) Achievements at CIRAD/INRA of triploid breeding of citrus with the 2x × 4x strategy Stone Fruits (Prunus) Peach Apricot Almond Plum Cherry Peach from Pompeii Stoneless Plum (Callahan Dardick Scorza JASH 1934 2009) Synteny in Prunus & Malus Prunus Malus Prunus Malus Prunus Malus Pome Fruits (Apple, Pear) Barrel Sprayer History of the PRI Program Dan Dayton Jules Janick Ed Williams Fred Hough Ralph Shay Sources of Scab-resistance Sources of Resistance Dominant Resistance M. atrosanguinea (2 genes) M. baccata jackii Dolgo M. floribunda 821 Geneva Jonsib M. micromalus M. prunifolia 19651 M. prunifolia microcarpa 782-26 M. prunifolia xanthocarpa 591-25 Morton Arboretum 4, 8, 16, 1255 Russian 12740-7A (2+ genes) Multigenic Resistance Antonovka M. baccata (selected seedlings) M. sargentii 843 M. sieboldii 2972-22 M. toringo 852 M. zumi calocarpa Defined Gene Pools Vf Vm Vr Vbj Vb Va M. floribunda M. micromalus (pit) Russian M. baccata jackii Hansens’s baccata #2 Antonovka (pit) Transferring Vf by Backcrossing PRI Releases Prima Priscilla Sir Prize Jonafree Redfree Dayton Williams’ Pride Enterprise Pristine GoldRush Scarlett O’Hara Pixie Crunch Sundance CrimsonCrisp 1970 1972 1975 1979 1981 1988 1988 1993 1993 1993 2000 2002 2002 2005 Joint Releases Viking (Wisc.) Priam (France) McShay (Ore.) Primiera (Italy) Nambu (Japan) Constance (Ger.) Primivera (Can.) Juliet (France) 1969 1974 1988 1995 1994 1995 1996 1999 Prima Redfree Sundance Pristine GoldRush Pixie Crunch Williams’ Pride Enterprise CrimsonCrisp Races of Venturia inaequalis Race Source 1 Worldwide 2 South Dakota, USA 3 4 5 6 7 Susceptible Material Most of the world’s cultivars M. baccata, ‘Dolgo’, ‘Alexis’, ‘Bittercrab’ segregates of R12740-7A, ‘Geneva’ Nova Scotia, Canada ‘Geneva’ Lafayette, IN, USA Segregates of R12740-7A Norwich, England Micromalus pit type resistance, M. atrosangunia 804 Ahrensburg, Germany ‘Prima’ (Vf cultivars) but not Evereste’ M. × ‘Perpetu’ and M. floribunda 821 England and Europe M. floribunda 821 Loquat (Eriobotrya japonica) Subtropical Pome Fruit Triploid Loquat Seedling ploidy Diploid Triploid Tetraploid Pentaploid Mixoploid No. seedlings 44,828 225 50 10 26 % 99.31 0.50 0.11 0.02 0.06 Seeded diploid loquat as compared to seedless triploid (courtesy of G. Liang) Molecular linkage map obtained from ‘Algerie’ and ‘Zaozhong-6’ Papaya Resistant to PRSV (D. Gonsalves) Transgenic ‘SunUp’-Hawaii (CP/CP) Transgenic ‘Rainbow’-Hawaii (CP/-) Transgenic ‘Khaknuan’ Thailand Tolerant ‘Thapra 2’ Thailand 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 Future Advances: 1. Search for specific mutations (parthenocarpy and seedlessness, breakdown of dioecy, loss of compatibility, high sugar, loss of astringency) 2. Use of interspecific crosses to create new fruits (Citrus, Prunus, Rubus) 3. Selection of underexploited germplasm (pitaya) 4. Induction of seedlessness 5. Transfer of genetic resistance to diseases and pests 6. Increased use of biotechnology (marker assisted selection, new traits, early flowering) 7. Emphasis on consumer satisfaction