Apple History—food of the gods, love, and good health Concepts

• Self-incompatible & Orchard Design • Cloning • Grafting • Germ plasm • New York State Agricultural Research Station – , grapes, cherries, hops

Kazakhstan is the Center of Diversity and Origin for Modern Apples Legend of James Chapman (a.k.a. ) • Credited with beginning apple nurseries across the Midwest (1802-1844)

• Homestead law to plant 50 apple trees on newly acquired property

• Apples were grown to make hard , not for eating Legend of John Chapman

• Preacher—Swedenborgism – The New Church

• Generous

• Vegetarian—practiced “humane” treatment to animals

• Unmarried, but he tried to cultivate a wife North American Apples Hardly an Aphrodisiac

Small, hard, green, & strongly astringent Chapman’s Great Selection Experiment

of apples discovered in North America – —Iowa – —Ohio – and Rome Beauty—New York Modern Apples = Golden Delicious x Jonathan

NY NZ GB Japan NY NY

• Cross-pollination of apple varieties • Selection among thousands of seedlings • Clonal propagation of trees onto good rootstocks

was indeed named after Cortland Co.— developed in Geneva, NY (1898) Natural populations in Kazakhstan hold the genes for improving modern apples Kazakhstan Apples

• Center of Genetic Diversity – Important breeding stock – Great variation in disease resistance, flavor, color, production • 2000-6000 B.C. spread of apples along trade routes to Middle East and southern Europe – Dessert Fruit, Aphrodisiac • Cloning techniques develop (i.e., grafting) Grafting—Scion of Desired

• Connect vascular cambium of scion with cambium of rootstock • Seal with wax & tape

University of Massachuesetts Grafting

• Root stock – Rate of growth determines size of tree • Dwarf (M9), semi-dwarf (M7, MM106), or standard (MM111) • M, Mailing England • MM, Mailing-Merton • G, Geneva NY

– Selection for • Disease resistance (fire blight & aphids) • Drought & flooding • Cold tolerance Healthy Living by Eating Apples

• Good sources of Fiber and Vitamin C • Phytonutrients – Flavonoids-anthocynanins-antioxidants – Anti-cancer compounds – Heart healthy compounds – Cholesterol lowering compounds On Trial in Geneva New York New York Agriculture Experimental Research Station in Geneva, NY

World’s Largest Germ Plasm Collection of Apples Apples credited to Geneva--Cortland Another invention from Geneva, NY What is it? Orchard Design

• Understanding apple incompatibility – Gametophytic system • Blooming time – Early, Mid, or Late Season Flowering • Pollinator Supplementation – Honey Bees—rental hives – Mason Bees, Bumble bees, orchard bees—habitat improvement • Environmental conditions – Wind, temperature extremes, nitrogen fertilizer Gametophytic Self- Incompatibility S S • “S” locus 9 1 – Hyper-variable • Crossing between trees of the same cultivar are incompatible • Diploid Genotypes

– S1S9 X S1S9; incompatible • Pollen tubes stopped shortly after entering stigma • Protein-protein interactions

– S1S9 X S2S9; partial compatible – S1S9 X S3S28; fully compatible

S1S9 Gametophytic Self- Incompatibility S S • “S” locus 3 28 – Hyper-variable • Diploid Genotypes

– S1S9 X S3S28; fully compatible – Pollen may possess an allele different from what is represented in the diploid, female parent.

S1S9 S-Locus Genotypes for some Apple Cultivars

—S10S28

—S2S5

• Ginger Gold—S3S28

• Golden Delicious—S2S3

—S3S23

—S2S24

• Jonathan—S7S9

—S1S3 Which apple cultivar appears to be

• Pink Lady—S2S23 the best pollinator of all the rest?

S-Locus Genotypes for some Apple Cultivars

• Empire—S10S28

• Gala—S2S5

• Ginger Gold—S3S28

• Golden Delicious—S2S3

• Granny Smith—S3S23

• Honeycrisp—S2S24

• Jonathan—S7S9

• Northern Spy—S1S3

• Pink Lady—S2S23 Which apple cultivar appears to be the best pollinator of all the rest?

S-Locus Genotypes & Bloom Times

• Empire—S10S28 • Early—Mid • Gala—S2S5 • Mid • Ginger Gold—S3S28 • Mid • Golden Delicious— • Mid S2S3 • Mid-Late • Granny Smith—S3S23 • Mid-Late • Honeycrisp—S2S24 • Early • Jonathan—S7S9 • Late • Northern Spy—S1S3 • Mid • Pink Lady—S2S23

S-Locus Genotypes & Bloom Times

• Empire—S10S28 • Early—Mid

• Gala—S2S5 • Mid

• Ginger Gold—S3S28 • Mid

• Golden Delicious—S2S3 • Mid

• Granny Smith—S3S23 • Mid-Late

• Honeycrisp—S2S24 • Mid-Late

• Jonathan—S7S9 • Early

• Northern Spy—S1S3 • Late

• Pink Lady—S2S23 • Mid

Apple growers must design orchards maximize compatibility & flowering times.

Sample Pollination Chart from Kelly Orchards Why are Gravestein, Jonagold, and King poor pollinators for all apples?

These are Triploid apples—pollen sterile, but require pollination for fruit-set.