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PBA workshop, MSU JUNE’ 10 1 QTL Discovery & Perspectives of use of markers in fruit breeding programs

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Page 1: 27 qtl discovery and-perspectives_e. van de weg

PBA workshop MSU JUNErsquo 10 1

QTL Discovery

amp Perspectives of use of

markers in fruit breeding programs

Content

Introduction

QTL-Discovery

Perspectives for use

Example Nectria (Canker) resistance

Summary

PBA workshop MSU JUNErsquo 10 2

Introduction

QTL discovery

Assessment of correlation trait harr molecular marker

Initial step towards Marker Assisted Breeding Discovery Validation High Throughput markers Logistics

Validation

Reproducibility on multiple families pedigrees

Mode of action

ndash Additive Dominance

ndash Gene x Gene interactions Independent action

Allelic diversity

Relative importance

PBA workshop MSU JUNErsquo 10 3

R R R S S S

Introduction

QTL Discovery

Assessment of correlation trait harr molecular marker

Requirements

1 Trait of interest

2 Phenotypic data

3 Segregating germplasm

4 Marker data genome-wide

5 QTL-mapping software

PBA workshop MSU JUNErsquo 10 4

QTL-Discovery in FruitBreedomics - Peach

1 Fruit related Traits

PBA workshop MSU JUNErsquo 10 5

ndash Maturity time

ndash Fruit Size

ndash Flesh color

ndash Firmsoft melting

ndash Flesh adherence

ndash Titrable acidity

ndash Total sugar

ndash Flowering time

ndash Flowering intensity

ndash White anthers

ndash Monilinia resistance

2 Phenotypic data

Historic

bull EU-Projects

bull National projects

New - FruitBreedomics

3 Segregating germplasm 18 families plusmn1500 individuals

5 breeding programs 2 countries

4 Marker Data 9K array

QTL-Discovery in FruitBreedomics - apple

1 Fruit Quality Traits gt 10

PBA workshop MSU JUNErsquo 10 6

Fruit Quality ndash Taste

bull Firmness bull Juiciness

bull Crispness bull Total Sugar bull Acidity

ndash Storability ndash Shelf life ndash Harvest date ndash Fruit size ndash Skin Color Resistance to Nectria

2 Phenotypic data

HiDRAS

bull 3 Years Multiple time moments

ndash Harvest Storage 2 amp 4M

ndash Shelf Life 2wks - 2M storage

National projects

3 Segregating germplasm 24 families plusmn1600 individuals

6 breeding programs 6 countries

4 Marker Data

Discovery Marker data

20K SNP array

New available 2013 Jan 16

First marker data Jan 18

Great performance

Some SSR

previous projects

2013 Exciting year

PBA workshop MSU JUNErsquo 10 7

Perspectives

PBA workshop MSU JUNErsquo 10 8

Great perspectives bull Many QTL discovered in EU-HiDRAS

bull Large intervals due to low marker density ( individuals high)

25

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Validation amp Marker development

On-going within FruitBreedomics

Development of SNP markers (WP1)

Firmness amp Storability

2 genes involved in ethylene pathway

Relevance confirmed in multiple studies

Already in use for MAB

bull WSU Russian programs

PBA workshop MSU JUNErsquo 10 9

Validation amp Marker development RosBREED

RosBREED project

US USDA-SCRI

Multi State International

Apple Peach Cherry Strawberry

wwwRosBREEDorg

Validation apple

Acidity

bull 2 loci including Ma locus

Crispness Juiciness (one LG)

Skin colour Rf locus

Bitter pit 3 loci

and others

Validation Peach

Acidity

Sweetness

Maturity date

Fruit size

and others

Perspectives on use in breeding

Discovery great potential

Validation completed for 1st genes

Marker Development started

Logistics started

Enhancement through intercontinental collaborations

RosBREED

Plant amp Food Research

PBA workshop MSU JUNErsquo 10 11

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

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45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

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25

30

35

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45

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55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

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30

35

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45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 2: 27 qtl discovery and-perspectives_e. van de weg

Content

Introduction

QTL-Discovery

Perspectives for use

Example Nectria (Canker) resistance

Summary

PBA workshop MSU JUNErsquo 10 2

Introduction

QTL discovery

Assessment of correlation trait harr molecular marker

Initial step towards Marker Assisted Breeding Discovery Validation High Throughput markers Logistics

Validation

Reproducibility on multiple families pedigrees

Mode of action

ndash Additive Dominance

ndash Gene x Gene interactions Independent action

Allelic diversity

Relative importance

PBA workshop MSU JUNErsquo 10 3

R R R S S S

Introduction

QTL Discovery

Assessment of correlation trait harr molecular marker

Requirements

1 Trait of interest

2 Phenotypic data

3 Segregating germplasm

4 Marker data genome-wide

5 QTL-mapping software

PBA workshop MSU JUNErsquo 10 4

QTL-Discovery in FruitBreedomics - Peach

1 Fruit related Traits

PBA workshop MSU JUNErsquo 10 5

ndash Maturity time

ndash Fruit Size

ndash Flesh color

ndash Firmsoft melting

ndash Flesh adherence

ndash Titrable acidity

ndash Total sugar

ndash Flowering time

ndash Flowering intensity

ndash White anthers

ndash Monilinia resistance

2 Phenotypic data

Historic

bull EU-Projects

bull National projects

New - FruitBreedomics

3 Segregating germplasm 18 families plusmn1500 individuals

5 breeding programs 2 countries

4 Marker Data 9K array

QTL-Discovery in FruitBreedomics - apple

1 Fruit Quality Traits gt 10

PBA workshop MSU JUNErsquo 10 6

Fruit Quality ndash Taste

bull Firmness bull Juiciness

bull Crispness bull Total Sugar bull Acidity

ndash Storability ndash Shelf life ndash Harvest date ndash Fruit size ndash Skin Color Resistance to Nectria

2 Phenotypic data

HiDRAS

bull 3 Years Multiple time moments

ndash Harvest Storage 2 amp 4M

ndash Shelf Life 2wks - 2M storage

National projects

3 Segregating germplasm 24 families plusmn1600 individuals

6 breeding programs 6 countries

4 Marker Data

Discovery Marker data

20K SNP array

New available 2013 Jan 16

First marker data Jan 18

Great performance

Some SSR

previous projects

2013 Exciting year

PBA workshop MSU JUNErsquo 10 7

Perspectives

PBA workshop MSU JUNErsquo 10 8

Great perspectives bull Many QTL discovered in EU-HiDRAS

bull Large intervals due to low marker density ( individuals high)

25

56

77

00

02

04

1

1

14

24

51

67

00

02

04

2

19

24

43

76

00

02

04

3

2

18

32

55

73

00

02

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4

8

14

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34

4547

69

90

100

10700

02

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5

9

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78

00

02

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26

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24

42

59

69

00

02

04

06

9

1314

40

50

64

81

104

00

02

04

10

13

29

45

51

61

76

00

02

04

12

3

18

30

45

64

89

00

02

04

13

8

23

37

77

110

116

00

02

04

15

8

14

30

47

76

00

02

04

16

0

12

2325

35

55

91

100

00

02

04

17

Validation amp Marker development

On-going within FruitBreedomics

Development of SNP markers (WP1)

Firmness amp Storability

2 genes involved in ethylene pathway

Relevance confirmed in multiple studies

Already in use for MAB

bull WSU Russian programs

PBA workshop MSU JUNErsquo 10 9

Validation amp Marker development RosBREED

RosBREED project

US USDA-SCRI

Multi State International

Apple Peach Cherry Strawberry

wwwRosBREEDorg

Validation apple

Acidity

bull 2 loci including Ma locus

Crispness Juiciness (one LG)

Skin colour Rf locus

Bitter pit 3 loci

and others

Validation Peach

Acidity

Sweetness

Maturity date

Fruit size

and others

Perspectives on use in breeding

Discovery great potential

Validation completed for 1st genes

Marker Development started

Logistics started

Enhancement through intercontinental collaborations

RosBREED

Plant amp Food Research

PBA workshop MSU JUNErsquo 10 11

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 3: 27 qtl discovery and-perspectives_e. van de weg

Introduction

QTL discovery

Assessment of correlation trait harr molecular marker

Initial step towards Marker Assisted Breeding Discovery Validation High Throughput markers Logistics

Validation

Reproducibility on multiple families pedigrees

Mode of action

ndash Additive Dominance

ndash Gene x Gene interactions Independent action

Allelic diversity

Relative importance

PBA workshop MSU JUNErsquo 10 3

R R R S S S

Introduction

QTL Discovery

Assessment of correlation trait harr molecular marker

Requirements

1 Trait of interest

2 Phenotypic data

3 Segregating germplasm

4 Marker data genome-wide

5 QTL-mapping software

PBA workshop MSU JUNErsquo 10 4

QTL-Discovery in FruitBreedomics - Peach

1 Fruit related Traits

PBA workshop MSU JUNErsquo 10 5

ndash Maturity time

ndash Fruit Size

ndash Flesh color

ndash Firmsoft melting

ndash Flesh adherence

ndash Titrable acidity

ndash Total sugar

ndash Flowering time

ndash Flowering intensity

ndash White anthers

ndash Monilinia resistance

2 Phenotypic data

Historic

bull EU-Projects

bull National projects

New - FruitBreedomics

3 Segregating germplasm 18 families plusmn1500 individuals

5 breeding programs 2 countries

4 Marker Data 9K array

QTL-Discovery in FruitBreedomics - apple

1 Fruit Quality Traits gt 10

PBA workshop MSU JUNErsquo 10 6

Fruit Quality ndash Taste

bull Firmness bull Juiciness

bull Crispness bull Total Sugar bull Acidity

ndash Storability ndash Shelf life ndash Harvest date ndash Fruit size ndash Skin Color Resistance to Nectria

2 Phenotypic data

HiDRAS

bull 3 Years Multiple time moments

ndash Harvest Storage 2 amp 4M

ndash Shelf Life 2wks - 2M storage

National projects

3 Segregating germplasm 24 families plusmn1600 individuals

6 breeding programs 6 countries

4 Marker Data

Discovery Marker data

20K SNP array

New available 2013 Jan 16

First marker data Jan 18

Great performance

Some SSR

previous projects

2013 Exciting year

PBA workshop MSU JUNErsquo 10 7

Perspectives

PBA workshop MSU JUNErsquo 10 8

Great perspectives bull Many QTL discovered in EU-HiDRAS

bull Large intervals due to low marker density ( individuals high)

25

56

77

00

02

04

1

1

14

24

51

67

00

02

04

2

19

24

43

76

00

02

04

3

2

18

32

55

73

00

02

04

4

8

14

20

34

4547

69

90

100

10700

02

04

5

9

40

78

00

02

04 6

26

35

00

02

04

7

18

33

52

62

00

02

04

8

24

42

59

69

00

02

04

06

9

1314

40

50

64

81

104

00

02

04

10

13

29

45

51

61

76

00

02

04

12

3

18

30

45

64

89

00

02

04

13

8

23

37

77

110

116

00

02

04

15

8

14

30

47

76

00

02

04

16

0

12

2325

35

55

91

100

00

02

04

17

Validation amp Marker development

On-going within FruitBreedomics

Development of SNP markers (WP1)

Firmness amp Storability

2 genes involved in ethylene pathway

Relevance confirmed in multiple studies

Already in use for MAB

bull WSU Russian programs

PBA workshop MSU JUNErsquo 10 9

Validation amp Marker development RosBREED

RosBREED project

US USDA-SCRI

Multi State International

Apple Peach Cherry Strawberry

wwwRosBREEDorg

Validation apple

Acidity

bull 2 loci including Ma locus

Crispness Juiciness (one LG)

Skin colour Rf locus

Bitter pit 3 loci

and others

Validation Peach

Acidity

Sweetness

Maturity date

Fruit size

and others

Perspectives on use in breeding

Discovery great potential

Validation completed for 1st genes

Marker Development started

Logistics started

Enhancement through intercontinental collaborations

RosBREED

Plant amp Food Research

PBA workshop MSU JUNErsquo 10 11

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 4: 27 qtl discovery and-perspectives_e. van de weg

Introduction

QTL Discovery

Assessment of correlation trait harr molecular marker

Requirements

1 Trait of interest

2 Phenotypic data

3 Segregating germplasm

4 Marker data genome-wide

5 QTL-mapping software

PBA workshop MSU JUNErsquo 10 4

QTL-Discovery in FruitBreedomics - Peach

1 Fruit related Traits

PBA workshop MSU JUNErsquo 10 5

ndash Maturity time

ndash Fruit Size

ndash Flesh color

ndash Firmsoft melting

ndash Flesh adherence

ndash Titrable acidity

ndash Total sugar

ndash Flowering time

ndash Flowering intensity

ndash White anthers

ndash Monilinia resistance

2 Phenotypic data

Historic

bull EU-Projects

bull National projects

New - FruitBreedomics

3 Segregating germplasm 18 families plusmn1500 individuals

5 breeding programs 2 countries

4 Marker Data 9K array

QTL-Discovery in FruitBreedomics - apple

1 Fruit Quality Traits gt 10

PBA workshop MSU JUNErsquo 10 6

Fruit Quality ndash Taste

bull Firmness bull Juiciness

bull Crispness bull Total Sugar bull Acidity

ndash Storability ndash Shelf life ndash Harvest date ndash Fruit size ndash Skin Color Resistance to Nectria

2 Phenotypic data

HiDRAS

bull 3 Years Multiple time moments

ndash Harvest Storage 2 amp 4M

ndash Shelf Life 2wks - 2M storage

National projects

3 Segregating germplasm 24 families plusmn1600 individuals

6 breeding programs 6 countries

4 Marker Data

Discovery Marker data

20K SNP array

New available 2013 Jan 16

First marker data Jan 18

Great performance

Some SSR

previous projects

2013 Exciting year

PBA workshop MSU JUNErsquo 10 7

Perspectives

PBA workshop MSU JUNErsquo 10 8

Great perspectives bull Many QTL discovered in EU-HiDRAS

bull Large intervals due to low marker density ( individuals high)

25

56

77

00

02

04

1

1

14

24

51

67

00

02

04

2

19

24

43

76

00

02

04

3

2

18

32

55

73

00

02

04

4

8

14

20

34

4547

69

90

100

10700

02

04

5

9

40

78

00

02

04 6

26

35

00

02

04

7

18

33

52

62

00

02

04

8

24

42

59

69

00

02

04

06

9

1314

40

50

64

81

104

00

02

04

10

13

29

45

51

61

76

00

02

04

12

3

18

30

45

64

89

00

02

04

13

8

23

37

77

110

116

00

02

04

15

8

14

30

47

76

00

02

04

16

0

12

2325

35

55

91

100

00

02

04

17

Validation amp Marker development

On-going within FruitBreedomics

Development of SNP markers (WP1)

Firmness amp Storability

2 genes involved in ethylene pathway

Relevance confirmed in multiple studies

Already in use for MAB

bull WSU Russian programs

PBA workshop MSU JUNErsquo 10 9

Validation amp Marker development RosBREED

RosBREED project

US USDA-SCRI

Multi State International

Apple Peach Cherry Strawberry

wwwRosBREEDorg

Validation apple

Acidity

bull 2 loci including Ma locus

Crispness Juiciness (one LG)

Skin colour Rf locus

Bitter pit 3 loci

and others

Validation Peach

Acidity

Sweetness

Maturity date

Fruit size

and others

Perspectives on use in breeding

Discovery great potential

Validation completed for 1st genes

Marker Development started

Logistics started

Enhancement through intercontinental collaborations

RosBREED

Plant amp Food Research

PBA workshop MSU JUNErsquo 10 11

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 5: 27 qtl discovery and-perspectives_e. van de weg

QTL-Discovery in FruitBreedomics - Peach

1 Fruit related Traits

PBA workshop MSU JUNErsquo 10 5

ndash Maturity time

ndash Fruit Size

ndash Flesh color

ndash Firmsoft melting

ndash Flesh adherence

ndash Titrable acidity

ndash Total sugar

ndash Flowering time

ndash Flowering intensity

ndash White anthers

ndash Monilinia resistance

2 Phenotypic data

Historic

bull EU-Projects

bull National projects

New - FruitBreedomics

3 Segregating germplasm 18 families plusmn1500 individuals

5 breeding programs 2 countries

4 Marker Data 9K array

QTL-Discovery in FruitBreedomics - apple

1 Fruit Quality Traits gt 10

PBA workshop MSU JUNErsquo 10 6

Fruit Quality ndash Taste

bull Firmness bull Juiciness

bull Crispness bull Total Sugar bull Acidity

ndash Storability ndash Shelf life ndash Harvest date ndash Fruit size ndash Skin Color Resistance to Nectria

2 Phenotypic data

HiDRAS

bull 3 Years Multiple time moments

ndash Harvest Storage 2 amp 4M

ndash Shelf Life 2wks - 2M storage

National projects

3 Segregating germplasm 24 families plusmn1600 individuals

6 breeding programs 6 countries

4 Marker Data

Discovery Marker data

20K SNP array

New available 2013 Jan 16

First marker data Jan 18

Great performance

Some SSR

previous projects

2013 Exciting year

PBA workshop MSU JUNErsquo 10 7

Perspectives

PBA workshop MSU JUNErsquo 10 8

Great perspectives bull Many QTL discovered in EU-HiDRAS

bull Large intervals due to low marker density ( individuals high)

25

56

77

00

02

04

1

1

14

24

51

67

00

02

04

2

19

24

43

76

00

02

04

3

2

18

32

55

73

00

02

04

4

8

14

20

34

4547

69

90

100

10700

02

04

5

9

40

78

00

02

04 6

26

35

00

02

04

7

18

33

52

62

00

02

04

8

24

42

59

69

00

02

04

06

9

1314

40

50

64

81

104

00

02

04

10

13

29

45

51

61

76

00

02

04

12

3

18

30

45

64

89

00

02

04

13

8

23

37

77

110

116

00

02

04

15

8

14

30

47

76

00

02

04

16

0

12

2325

35

55

91

100

00

02

04

17

Validation amp Marker development

On-going within FruitBreedomics

Development of SNP markers (WP1)

Firmness amp Storability

2 genes involved in ethylene pathway

Relevance confirmed in multiple studies

Already in use for MAB

bull WSU Russian programs

PBA workshop MSU JUNErsquo 10 9

Validation amp Marker development RosBREED

RosBREED project

US USDA-SCRI

Multi State International

Apple Peach Cherry Strawberry

wwwRosBREEDorg

Validation apple

Acidity

bull 2 loci including Ma locus

Crispness Juiciness (one LG)

Skin colour Rf locus

Bitter pit 3 loci

and others

Validation Peach

Acidity

Sweetness

Maturity date

Fruit size

and others

Perspectives on use in breeding

Discovery great potential

Validation completed for 1st genes

Marker Development started

Logistics started

Enhancement through intercontinental collaborations

RosBREED

Plant amp Food Research

PBA workshop MSU JUNErsquo 10 11

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 6: 27 qtl discovery and-perspectives_e. van de weg

QTL-Discovery in FruitBreedomics - apple

1 Fruit Quality Traits gt 10

PBA workshop MSU JUNErsquo 10 6

Fruit Quality ndash Taste

bull Firmness bull Juiciness

bull Crispness bull Total Sugar bull Acidity

ndash Storability ndash Shelf life ndash Harvest date ndash Fruit size ndash Skin Color Resistance to Nectria

2 Phenotypic data

HiDRAS

bull 3 Years Multiple time moments

ndash Harvest Storage 2 amp 4M

ndash Shelf Life 2wks - 2M storage

National projects

3 Segregating germplasm 24 families plusmn1600 individuals

6 breeding programs 6 countries

4 Marker Data

Discovery Marker data

20K SNP array

New available 2013 Jan 16

First marker data Jan 18

Great performance

Some SSR

previous projects

2013 Exciting year

PBA workshop MSU JUNErsquo 10 7

Perspectives

PBA workshop MSU JUNErsquo 10 8

Great perspectives bull Many QTL discovered in EU-HiDRAS

bull Large intervals due to low marker density ( individuals high)

25

56

77

00

02

04

1

1

14

24

51

67

00

02

04

2

19

24

43

76

00

02

04

3

2

18

32

55

73

00

02

04

4

8

14

20

34

4547

69

90

100

10700

02

04

5

9

40

78

00

02

04 6

26

35

00

02

04

7

18

33

52

62

00

02

04

8

24

42

59

69

00

02

04

06

9

1314

40

50

64

81

104

00

02

04

10

13

29

45

51

61

76

00

02

04

12

3

18

30

45

64

89

00

02

04

13

8

23

37

77

110

116

00

02

04

15

8

14

30

47

76

00

02

04

16

0

12

2325

35

55

91

100

00

02

04

17

Validation amp Marker development

On-going within FruitBreedomics

Development of SNP markers (WP1)

Firmness amp Storability

2 genes involved in ethylene pathway

Relevance confirmed in multiple studies

Already in use for MAB

bull WSU Russian programs

PBA workshop MSU JUNErsquo 10 9

Validation amp Marker development RosBREED

RosBREED project

US USDA-SCRI

Multi State International

Apple Peach Cherry Strawberry

wwwRosBREEDorg

Validation apple

Acidity

bull 2 loci including Ma locus

Crispness Juiciness (one LG)

Skin colour Rf locus

Bitter pit 3 loci

and others

Validation Peach

Acidity

Sweetness

Maturity date

Fruit size

and others

Perspectives on use in breeding

Discovery great potential

Validation completed for 1st genes

Marker Development started

Logistics started

Enhancement through intercontinental collaborations

RosBREED

Plant amp Food Research

PBA workshop MSU JUNErsquo 10 11

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 7: 27 qtl discovery and-perspectives_e. van de weg

Discovery Marker data

20K SNP array

New available 2013 Jan 16

First marker data Jan 18

Great performance

Some SSR

previous projects

2013 Exciting year

PBA workshop MSU JUNErsquo 10 7

Perspectives

PBA workshop MSU JUNErsquo 10 8

Great perspectives bull Many QTL discovered in EU-HiDRAS

bull Large intervals due to low marker density ( individuals high)

25

56

77

00

02

04

1

1

14

24

51

67

00

02

04

2

19

24

43

76

00

02

04

3

2

18

32

55

73

00

02

04

4

8

14

20

34

4547

69

90

100

10700

02

04

5

9

40

78

00

02

04 6

26

35

00

02

04

7

18

33

52

62

00

02

04

8

24

42

59

69

00

02

04

06

9

1314

40

50

64

81

104

00

02

04

10

13

29

45

51

61

76

00

02

04

12

3

18

30

45

64

89

00

02

04

13

8

23

37

77

110

116

00

02

04

15

8

14

30

47

76

00

02

04

16

0

12

2325

35

55

91

100

00

02

04

17

Validation amp Marker development

On-going within FruitBreedomics

Development of SNP markers (WP1)

Firmness amp Storability

2 genes involved in ethylene pathway

Relevance confirmed in multiple studies

Already in use for MAB

bull WSU Russian programs

PBA workshop MSU JUNErsquo 10 9

Validation amp Marker development RosBREED

RosBREED project

US USDA-SCRI

Multi State International

Apple Peach Cherry Strawberry

wwwRosBREEDorg

Validation apple

Acidity

bull 2 loci including Ma locus

Crispness Juiciness (one LG)

Skin colour Rf locus

Bitter pit 3 loci

and others

Validation Peach

Acidity

Sweetness

Maturity date

Fruit size

and others

Perspectives on use in breeding

Discovery great potential

Validation completed for 1st genes

Marker Development started

Logistics started

Enhancement through intercontinental collaborations

RosBREED

Plant amp Food Research

PBA workshop MSU JUNErsquo 10 11

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 8: 27 qtl discovery and-perspectives_e. van de weg

Perspectives

PBA workshop MSU JUNErsquo 10 8

Great perspectives bull Many QTL discovered in EU-HiDRAS

bull Large intervals due to low marker density ( individuals high)

25

56

77

00

02

04

1

1

14

24

51

67

00

02

04

2

19

24

43

76

00

02

04

3

2

18

32

55

73

00

02

04

4

8

14

20

34

4547

69

90

100

10700

02

04

5

9

40

78

00

02

04 6

26

35

00

02

04

7

18

33

52

62

00

02

04

8

24

42

59

69

00

02

04

06

9

1314

40

50

64

81

104

00

02

04

10

13

29

45

51

61

76

00

02

04

12

3

18

30

45

64

89

00

02

04

13

8

23

37

77

110

116

00

02

04

15

8

14

30

47

76

00

02

04

16

0

12

2325

35

55

91

100

00

02

04

17

Validation amp Marker development

On-going within FruitBreedomics

Development of SNP markers (WP1)

Firmness amp Storability

2 genes involved in ethylene pathway

Relevance confirmed in multiple studies

Already in use for MAB

bull WSU Russian programs

PBA workshop MSU JUNErsquo 10 9

Validation amp Marker development RosBREED

RosBREED project

US USDA-SCRI

Multi State International

Apple Peach Cherry Strawberry

wwwRosBREEDorg

Validation apple

Acidity

bull 2 loci including Ma locus

Crispness Juiciness (one LG)

Skin colour Rf locus

Bitter pit 3 loci

and others

Validation Peach

Acidity

Sweetness

Maturity date

Fruit size

and others

Perspectives on use in breeding

Discovery great potential

Validation completed for 1st genes

Marker Development started

Logistics started

Enhancement through intercontinental collaborations

RosBREED

Plant amp Food Research

PBA workshop MSU JUNErsquo 10 11

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 9: 27 qtl discovery and-perspectives_e. van de weg

Validation amp Marker development

On-going within FruitBreedomics

Development of SNP markers (WP1)

Firmness amp Storability

2 genes involved in ethylene pathway

Relevance confirmed in multiple studies

Already in use for MAB

bull WSU Russian programs

PBA workshop MSU JUNErsquo 10 9

Validation amp Marker development RosBREED

RosBREED project

US USDA-SCRI

Multi State International

Apple Peach Cherry Strawberry

wwwRosBREEDorg

Validation apple

Acidity

bull 2 loci including Ma locus

Crispness Juiciness (one LG)

Skin colour Rf locus

Bitter pit 3 loci

and others

Validation Peach

Acidity

Sweetness

Maturity date

Fruit size

and others

Perspectives on use in breeding

Discovery great potential

Validation completed for 1st genes

Marker Development started

Logistics started

Enhancement through intercontinental collaborations

RosBREED

Plant amp Food Research

PBA workshop MSU JUNErsquo 10 11

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 10: 27 qtl discovery and-perspectives_e. van de weg

Validation amp Marker development RosBREED

RosBREED project

US USDA-SCRI

Multi State International

Apple Peach Cherry Strawberry

wwwRosBREEDorg

Validation apple

Acidity

bull 2 loci including Ma locus

Crispness Juiciness (one LG)

Skin colour Rf locus

Bitter pit 3 loci

and others

Validation Peach

Acidity

Sweetness

Maturity date

Fruit size

and others

Perspectives on use in breeding

Discovery great potential

Validation completed for 1st genes

Marker Development started

Logistics started

Enhancement through intercontinental collaborations

RosBREED

Plant amp Food Research

PBA workshop MSU JUNErsquo 10 11

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 11: 27 qtl discovery and-perspectives_e. van de weg

Perspectives on use in breeding

Discovery great potential

Validation completed for 1st genes

Marker Development started

Logistics started

Enhancement through intercontinental collaborations

RosBREED

Plant amp Food Research

PBA workshop MSU JUNErsquo 10 11

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 12: 27 qtl discovery and-perspectives_e. van de weg

Nectria Canker

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 13: 27 qtl discovery and-perspectives_e. van de weg

Introduction on Nectria

PBA workshop MSU JUNErsquo 10 13

Causes canker in apple

Problem in NW-Europe (Humid climates)

Many (popular) cvs extremely susceptible

Braeburn Cox Gala Kanzi Rubens Prima

Fungicides for Scab act also to Nectria

Problems with Nectria may increase

Scab resistance

Susceptible parents

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 14: 27 qtl discovery and-perspectives_e. van de weg

Introduction to Nectria

Breeding for resistance is hard

R ndash highly quantitative

Only few cultivars show reasonable resistance

bull Jonathan Golden Delicious Lombartrsquos Calvillle Santana

Resistance tests are laborious and time-lasting

Availability of markers highly useful

Difficult to develop

PBA workshop MSU JUNErsquo 10 14

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

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45

50

55

60

65

70

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105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

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5

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65

70

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105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

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55

60

65

70

75

80

85

90

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105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

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65

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80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 15: 27 qtl discovery and-perspectives_e. van de weg

Discovery

Germplasm

Jonathan x Prima

bull Jonathan cv with highest level of resistance

bull Prima highly susceptible chopped down in biological production

PBA workshop MSU JUNErsquo 10 15

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 16: 27 qtl discovery and-perspectives_e. van de weg

Discovery National Project

Germplasm Jonathan x Prima

PBA workshop MSU JUNErsquo 10 16

Phenotyping

Artificial inoculation (1994)

Greenhouse no 2nd infection

Colonization rate Length of cankers in time (1996)

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 17: 27 qtl discovery and-perspectives_e. van de weg

Phenotypic data Frequency Distribution

Size of cankers (in mm)

p

roge

ny

0

5

10

15

20

25

30

35

56

4

8

12

16

20

24

28

32

36

40

44

48

52

60

64

68

72

76

1

28

25

16

12

5 4

2 1

13

30

18

8 6

3

1

4 5

3

1 1 1 1 1 1 1

bull Prima Large experimental variation

Jonathan x Prima

Jonathan

Prima

bull JP-Population sub-groups

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 18: 27 qtl discovery and-perspectives_e. van de weg

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5-

438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638CH05g08-175178+- 648

E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M47-128-10- 00

E31M56-185-4+ 65E33M47-113-15-Fd+ 77

E32M58-352-1- 118

E35M47-253-9-Fd- 149E35M61-163-6+ 154

E33M47-290-3+ 197

E35M53-191-4+ 224

MC036c- 267

E31M52-140-14- 490E31M52-139-9- 492E34M54-142-7- 493

E31M52-178-12- 505

E33M60-92-20+ 705

E35M54-347-2 - 00E35M50-215-5 - 28E35M51-154-16-Pf-fD - 34

E33M60-172-11-fD + 75

E34M60-99-12 + 233E34M50-180-7 - 250

E34M48-308-28-fD-309 + 319E34M60-264-4 - 320E34M47-170-6 + 323E34M60-108-11 - E32M58-106-8 -

337

E34M55-278-6-Pf-272 - 412

E35M53-120-7 + 491

E32M52-171-7 -636

E31M48-173-16-Fd+ 00

CH04e02-155148-+ 34

E35M51-198-14- 189E34M47-137-9- 190

E34M50-122-14- 191E31M48-127-20- 201E34M55-317-5- 204

GD162-223247-+ 255E35M52-292-pF-3- 262

CH02h11a-104120-+ 292

E32M50-395-2+ 507

E35M54-206-5- 614

E34M51-250-5- 667

CH04e02-155163 -+00E34M51-100-16 -05E34M59-291-3-Pf +08E33M60-286-6 +15MC036a -23E33M51-207-6-Pf -48

E33M60-148-13 -239

CH01d03-138158 -266E32M48-439-1 -313

CH02h11a-124130 +-314E35M51-88-18 -315E33M51-225-4-Pf-fD +322

E35M61-256-4 -454E34M51-278-3 -473

CH02c02b-112116 +-595

J3 P3 J4 P4

CH03g12z-170154+- 00

E35M50-137-12- 100E35M53-168-6- 122E34M49-199-3- 125

E35M52-133-7+ 265E33M47-97-16+ 274

E35M47-310-4-pF-Fd+ 288

E31M52-90-13-E35M51-305-pF-5- 438

E33M47-76-17- 440E31M52-90-18- 441

E35M47-200-12- 489

E34M59-310-2- 638

CH05g08-175178+- 648E34M51-122-13- 660

MC014d -00

CH03g12z-170162+-191E36M51-145-16-fD -197E36M51-397-3 -223

E35M52-81-11-Pf +334

E33M60-98-18 +494

E31M52-302-4-Pf +564

E35M47-361-1 +645E32M50-169-10 +664

E34M54-244-4 +818E34M51-133-11 +819E34M51-115-14 +824CH05g08-1750 -+839

CH02f06-158150-+ 00E31M48-357-4+ 30

E34M48-171-18-pF-Fd+E34M53-221-5+

36

E35M47-278-8+E35M47-224-10+ 92

E34M55-430-2+ 94

E35M49-163-11+ 342E35M50-210-6+ 369E33M60-321-4+ 370

E34M54-111-12+ 375E32M58-116-6+ 392E34M51-153-8+

E32M47-119-11+ 396

CH05e03-182161-+ 398MC029a- 479

CH03d01-1130-+ 482E34M48-262-10- 487E35M52-89-10- 501

MC064b- 518

E34M53-470-1 - 00

E33M60-96-19 -172E34M47-298-1 -174

E35M55-118-8 -232

E34M59-192-Pf-9 -272

CH05e03-182176+-279

CH03d01-113111-+362

E34M50-110-16 +415

E34M48-288-29 +444

J1 P1 J2 P2

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M53-169-7- 00

MC064a- 64

E36M51-118-19+ 124

E33M51-136-11-fD+ 176

E32M47-107-15+ 298

CH04e05-174201-+ 309E35M58-242-2+

E32M47-110-14+310

E32M48-400-3+ 324E32M48-151-8-Fd+ 330

E35M47-220-11-pF-Fd- 355

E34M56-220-4+ 588E32M50-400-1+ 604

E31M48-89-25+ 734

J7E35M48-81-12 +00

E34M49-96-6 -43MC064a -53E34M50-198-6 +73E31M48-365-3-fD -80E34M50-171-10 -88E35M58-91-9 +120E34M55-283-19 -149E36M51-206-8-Pf-fD +155

E31M48-350-5 +210

E34M47-213-3 +247

E31M48-187-14 +287E35M59-255-1 -299E35M48-226-4 +302E33M60-278-7 -313E35M48-131-9-Pf +349

E35M47-280-7 -394

CH04e05-174209 +-396

E32M58-470-12 +588

P7

E31M49-206-5-Fd- 00E34M56-230-2- 11E32M58-228-3+ 32

E34M60-206-8-Fd+ 53

Ly37c+ 136E31M52-465-1-pF+ 137

E35M54-61-15+ 148MC022a+ 154

E35M61-75-8+ 181

E32M48-109-13-Fd+E32M48-63-16-Fd+

E32M48-60-20+426

J8E34M53-97-14 + 00E34M48-218-31 + 26E34M50-175-9 - 45E33M51-67-18 - 51E34M59-203-8 - 55E34M56-228-3 - 67

E34M48-323-6-Pf - 133Ly37c - 160E31M52-232-8 - 167

MC022a - 203E34M48-318-7 - 206E34M57-157-10 - 233E35M58-132-6 - 248E34M57-179-9 - 265E35M53-177-5 - 288

P8

E34M57-190-8+ 00E34M50-115-15- 28

CH04e03-186196+- 56E34M49-86-7- 66

E31M48-483-1-Fd+ 72

CH03a04-100114+- 108

CH02b12-125127+- 125E32M48-130-11+ 126E35M61-308-2- 129

E34M48-154-19-pF- 132E34M55-59-18-pF+ 136

E31M48-185-15-Fd- 141E34M48-115-25- 143

E31M52-215-6Fd-E34M54-72-18+E34M59-96-14+

145

E35M47-158-14+ 146E33M60-304-5-E33M60-340-3+

147

E33M60-213-9+ 155E34M55-85-14-pF+ 160

E35M50-202-7- 168

GD103-114118+- 171E35M53-230-3- 280E35M49-227-8+ 345

E34M53-138-10+ 347E33M51-221-5+ 362

MC025a- 419E33M60-358-2- 447

CH05e06-0150+- 463E36M51-190-11- 468E31M49-385-2- 483E34M60-207-7- 496E35M59-177-6- 506

CH04e03-184204 +- 00E34M57-352-2 + 47E35M59-103-8 +54E35M54-115-9 -96E34M54-105-13 -E32M50-119-13 -E34M49-316-1 +

97

E32M58-98-9 -E32M58-207-13 -E31M52-376-3 -E33M60-104-16 -E33M60-178-10-fD-179 -

98

E33M51-202-7-Pf -99E35M54-69-13 +100E35M47-126-20 -104E35M49-124-14 -115E35M61-307-3 -124E33M60-131-14 -127

CH02b12-129141 -+

CH03a04-0106 +130

E34M48-550-1 +138E34M49-112-5 +207E34M48-440-3 +213E32M50-145-11 -253E34M59-251-6-Pf -307E32M52-375-2-Pf +355E33M51-127-14-Pf-fD -483E35M49-81-18 -501MC025a -515E34M51-103-15 +549E34M51-406-1 -601

CH05e06-0134 -633

E34M50-292-2 +678

E35M49-397-1 -867

E35M50-185-8 +890

E34M48-141-20- 00

E33M47-223-8- 42MC014c- 47

E35M51-158-15-Fd+ 99

CH03d07-206202- 102MC023a+ 119

E34M55-122-10+ 125

E34M48-220-12+E31M52-113-11+

256

E35M59-150-7+ 272

E34M57-300-3+ 413

CH05a05-198218+- 529

CH03c01-172174-+ 548E34M48-257-11+ 561

CH03d07-206218 -+00

E32M48-185-19 +125

E34M57-122-13-fD +151

E31M49-471-13 -203

CH03d12-124150 +-341

E34M57-293-4 +482E34M55-84-16-Pf +495

CH05a05-198218 -+613

CH03c01-172174 +-633

J5 P5 J6 P6

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E34M59-155-10-Fd- 00

E35M49-308-6- 68

E35M55-171-7+ 94E36M51-196-10- 115

E34M53-52-18- 235E32M52-433-1- 241

CH05c07-141149+- 283

E31M51-247-2- 431E35M47-322-3- 455

E36M51-480-1- pF- 464E34M48-83-24+ 466E35M58-174-4+E34M60-257-5+

467

E32M50-90-15- Fd+ 473

CH01h02b-236248+- 491E34M48-103-23- 521

J9-5

E35M61-92-7 + 00

E34M57-102-16 +22

E34M51-123-12-Fd + 76E34M47-267-2 -93

E34M57-111-15- fD -210

E33M60-357-1- fD -248E36M51-167-13-Pf -254

E35M49-155-12 -288E35M47-490-22 -307

CH05c07-141149 -+386

E34M53-150-9 +494E32M52-239-6

+504

E32M52-242-5 +565E32M48-132-10 +567E35M50-380-1 +579E35M47-480-21 +E31M48-132-19 -

582

E31M49-303-3 - 596E32M47-169-7 + 608E32M48-119-12 -618

CH01h02b-236246 +- 655

P9-3

E34M54-206-5-Fd+ 00E32M48-80-15+ 01

E34M50-156-11-Fd+ 06E34M53-120-11- 40

CH04c06z-157169-+ 46

CH02b07-202180-+ 57E34M47-100-13- 58

E34M59-90-15- 62E35M49-419-19- 70E35M47-106-24+

E35M58-113-7+89

CH02a08-151-+ 91E34M51-310-2+ 113

E36M51-102-21+ 118E35M59-220-3+ 128

E31M48-102-22-Fd+ 131

CH02a10-177- 132E32M47-228-1+ 151

E34M59-109-13- 152CH02c11-231237-+ 198

E35M53-357-2+

CH04g09-145-

320

COL-230232+- 475

E34M54-122-11+ 510

CH02b03b-9799t1=-+CH01f07a-194196-+ 559

GD100-236+ 618E35M58-251-1- 634

J10-4

CH04c06z-157169 +- 00E34M50-300-1 -

CH02b07-184202 +- 01

CH02a08-147151 +- 79

E31M48-66-26 -139E35M52-243-4 - 145

CH02a10-149155 +- 177E32M47-126-9 - 193E34M59-314-1-Pf - 195

CH02c11-231235 -+ 293E34M47-105-11 - 309

E32M47-171-6 - 352

CH03d11-121127+179 +- 353

E32M48-212-7 +409E35M52-224-5-Pf -426

CH04g09-153 +453

E35M48-159-7-Pf + 476

CH02b03b-9397 -+CH01f07a-194190 -+

520

COL-220234 +- 545

E34M54-82-17 - 610

E31M48-250-8 -676

P10-5

E33M60-270-8- 00

E35M47-177-13+ 53

E32M48-242-6+ 80E31M49-214-4- 128

CH03d08-228+- 136

CH03d02-203+- 137E35M52-117-8+ 140

CH04a12-180195-+ 253

E33M51-182-9- 302E35M54-67-14-E35M52-375-2-

311

E33M47-115-14- 338E35M52-171-6- 348E34M53-69-16- 355E34M53-360-2- 387

E34M59-68-17- 444

E34M53-107-13- Fd+ 514

E35M59-182-5+ 560

J11-5

E34M59-139-11- Pf +00

E34M54-165-8 +39

E34M50-147-12 +88E31M49-169-7 +89

E34M55-410-3- Pf +174

E34M57-77-19- fD -217

CH04a12-172176 +-286E32M48-287-5 -324E33M51-96-16 -325E32M47-115-13 -329

E32M48-339-4 -375

E34M56-187-5 -417E33M51-133-13- Pf -432E34M55-83-15 -451

E31M49-388-1 -555E34M57-148-11 -559E32M50-189-6 -575

E34M60-410-1 -628

E35M52-400-1 +687

E34M53-228-3 +716E34M55-91-13- Pf +733E34M54-97-14 -752

E35M59-187-4 -808E34M59-260-4 +836E31M48-338-6- fD -846E35M61-384-1 -852

P11-4

CH05d04-174180-+ 00MC032b- 23MC225a- 31

CH04g04-180168-+ 64

E35M48-291-3+ 99E34M51-145-9- Fd+ 100

E33M51-236-3+ 170E34M54-87-16+ 180

CH01g12-110186+- 193E31M52-292-2- 200

E31M52-290-5- pF-Fd- 204

CH04d02-154+- 263

E34M53-159-8+ 294

E35M59-76-9- 435

E34M48-173-17+ 482

J12-4

CH05d04-176186 -+00MC225a +03E32M48-58-17 +04MC032b +07E31M48-248-9 +15E33M47-143-11 -34

CH04g04-180172 -+62E32M58-83-11- fD +82E34M57-117-14- fD +134E35M48-196188-6- Pf -147E34M51-144-10- fD +149E34M60-79-13 -184

E31M48-119-21 -259

E34M53-417-19 - 297

E32M50-202-4 -557

E35M49-127-13 +633E35M47-127-19- fD +634E32M58-224-4 +659

P12-5

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

E31M48-150-18 -796

E32M58-316-2- 00

CH02g01-217+ 09E31M52-127-15- 28E31M52-127-10- 36E34M51-164-6- 43

E32M52-120-9- 133

E31M48-100-23+ 267E34M51-159-7+ 270

E33M47-140-12+ 316

E34M53-191-6 +00

E34M48-283-8 +52E32M58-92-10 +58E34M59-255-5-Pf +72

CH02g01-0226 -91

E34M60-272-3 -265

E35M54-137-8 -357

E31M48-165-17 +507

E35M50-219-4 -E31M49-139-8 -627

E35M54-207-6 -630

CH03h03-91103 +-634E33M47-348-2 -666

CH05f04-162158 +-E34M59-118-12 -

669

E35M58-134-5 -E31M52-158-8 -E33M47-165-10 -E34M48-419-4-fD -

671

E31M52-158-13 -677

E34M48-275-9- 00

E31M56-328-1- 39E32M47-209-3+ 50

E34M48-182-15-pF-Fd- 102E34M50-265-3- 110

E34M55-106-12- 207E31M52-75-14- 220

CH05g11-243233+- 227

CH04c07-112114-+ 246

CH05d03-158173+- 330E31M56-188-3+ 333

E34M57-250-7-Fd+ 362E34M60-290-2a-Fd+ 378

E34M54-340-19- 418E34M55-167-8+ 431E34M53-225-4- 436

E32M52-148-8- 469

E31M48-228-10 -00

E31M48-195-12 -109

CH04f06-175178 -+ 136E34M48-179-16-Pf +151E32M47-92-19 +161E33M51-112-15 +167E33M47-172-9 +197

E33M47-275-4 -273

CH05g11-2430 +-328

CH04c07-106108 -+338

CH05d03-152179 -+432E34M60-247-6 +449E34M49-170-4 -456E35M47-149-16-Pf-fD +473E34M55-166-9 +478

E35M58-95-8 -586

J13 P13 J14 P14

+-

+-

E34M50-177-8- 00

CH05c06-104118 163E35M47-287-6- 171

CH05e04a-159167 186

LY05b- 224

E32M52-264-4+ 341

E31M48-435-2- 420E32M47-129-8+ 466E34M47-198-4+ 467E31M52-52-20+E32M58-113-7+

517

E35M53-91-8+ 556E34M48-122-22+ 565E32M48-51-18+ 569

E32M52-114b-10-pF-E34M59-243-7-

570

E35M54-388-16- 571E35M61-218-5+ 573

CH05e04a-1590 -+00

CH05c06-104110 -+ 02E35M50-99-15 + 34

E34M57-292-5 + 104

E33M47-357-1 - 218

E31M49-81-9 + 260

CH05a04-163183 +- 278

E31M49-480-14 -480E31M51-257-1 +490E31M52-218-5 -562E34M54-200-6 -598E32M58-176-5-fD +615E34M50-101-17 -616E35M59-228-2 -E32M52-113-12-Pf -

637

E35M49-108-16 +658E32M52-330-3-Pf +659E34M51-276-4 -670E34M48-140-21 +E34M48-138-32 +674

E33M60-107-15-fD +679

E34M48-130-27 -790

J16 P16

Marker data

AFLP amp SSR

1998-2000

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 19: 27 qtl discovery and-perspectives_e. van de weg

QTL ndashmapping

PBA workshop MSU JUNErsquo 10 19

Effect QTL

Additive

Q1+Q2 -gt Cankers as small as for Jonathan

bull Prima 47 mm

bull Jonathan 23 mm

bull JxP_Q1Q2 23 mm

Highly effective

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

3 QTL identified

2x Jonathan 1x Prima

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 20: 27 qtl discovery and-perspectives_e. van de weg

Fruitbreedomics

PBA workshop MSU JUNErsquo 10 20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

P-Q3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q2 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

105

J-Q1

Position

J-Q1

bull No markers for distal part

bull Sub-optimum

J-Q2

bull 1 marker 35 cM

bull Position

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 21: 27 qtl discovery and-perspectives_e. van de weg

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

PBA workshop MSU JUNErsquo 10 21

J01 J02 J03 J04 J05 J06 J07 J08 J09 J10 J11 J12 J13 J14 J15 J16 J17

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

bull SNP-array plusmn8000 markers for JxP

bull plusmn3000 for Jonathan

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 22: 27 qtl discovery and-perspectives_e. van de weg

Fruitbreedomics

Progresses on these historic data

Fine-mapping

Increase marker density

Validation

Reproducibility 2nd mapping population

Extension

Stability over environments

bull Phenotype JxP in Sweden

Other parameters for resistance

bull Infection percentage

Larisa Gustavsson

Hilde Nybom

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23

Page 23: 27 qtl discovery and-perspectives_e. van de weg

Summary

Title QTL Discovery amp Perspectives of use of markers in fruit breeding programs

QTL Discovery 1st step to application

Discovery Validation High Throughput markers Logistics

Assessment of correlation trait harr molecular marker

Potential - couple of years

Apple gt 10 fruit quality traits + Nectria resistance

Peach gt Flowering + Fruit traits + Monilinia resistance

Potential - short term (from building on other projects)

Apple Fruit firmness (ACS ACO) Skin Color (Rf) Acidity (Ma) Bitter pit

Peach Sub-acidAcid Flatround (PeachNectarine Yellowwhite flesh)

Pipeline-Logistics under development

PBA workshop MSU JUNErsquo 10 23