Genomics of Environmental Stress Responses in Hardwoods€¦ · 10/5/2011 6 1000 Cankers Disease...

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"Genomics of Environmental Stress Responses in Hardwoods"

John CarlsonProfessor of Molecular GeneticsProfessor of Molecular Genetics

Director, Schatz Center for Tree Molecular GeneticsThe School of Forest Resources, Pennsylvania State University

Populations of forest trees represent nature’s long-term experiments in adaptation which are rich, largely untapped resources for evolutionary and ecological genomics.

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However, the sustainability of many of tree species and populations are threatened by forest health issues:

• introduced insects,

• exotic diseasesexotic diseases,

• invasive plants,

• climate change,

• fragmentation,

• poor management practices.

A forest health crisis is underwayA forest health crisis is underway.

Many important tree species and forest ecosystems are threatened by pests, diseases and climate change.

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Many Forest Tree Species Under Attack

• Castanea dentata

• Ulmus americana

• Pinus strobus

• Pinus contorta

• Abies balsamifera

• Tsuga canadensis, caroliniana

• Fraxinus spp.

• Juglans cinceraJuglans cincera

• Juglans nigra

• Fagus grandifolia

• Quercus spp.

Emerald Ash BorerAgrilus planipennis

• Invasive native to Asiaintroduced in 2002 onwood packing material

• Killing most ash speciesof all age classes

• Firewood key to spread

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Asian Longhorned BeetleAnoplophora glabripennis

• Invasive species introduced from Asiaon wood packing materials in 1998p g

• Declared eradicated from Chicago, ILon April 17, 2008

• New infestation discovered inWorcester, MA in August 2008

Chicago 1998

*

*

New York 1996

Toronto 2003

*New Jersey

2002 *

Massachusetts 2008

*• Present in MA at least 7 yrs

Hemlock Woolly AdelgidAldeges tsugae

• Introduced insect, native of Asia,is a serious pest of easternphemlock and Carolina hemlock

• In the United States since 1924

• Biological control agents havebeen released, with limitedsuccess to date.success to date.

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David Vance

European Wood WaspSirex noctilio

• Current infestation on Scots pineand red pine stands in NY andsurrounding statessurrounding states

• Infestation in other countries hascaused severe mortality in pineplantations

• Threat to Monterey, lodge‐pole,d j k & t thponderosa, jack & most southern

pines (especially loblolly)

• The susceptibility of other USconifers is not known

Sudden Oak DeathPhytophthora ramorum

Sudden Oak DeathPhytophthora ramorum

Invasive pathogen along the CA and Invasive pathogen along the CA and Edwin FlorenceOR coasts

Extensive mortality of tanoak, coast live oak, and CA black oak since 1995

Surveys have not found forest

OR coasts

Extensive mortality of tanoak, coast live oak, and CA black oak since 1995

Surveys have not found forestSurveys have not found forest infestations outside CA and ORSurveys have not found forest infestations outside CA and OR

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1000 Cankers DiseaseGeosmithia sp.

• New canker disease vectored bywalnut twig beetle Pityophthorus

Ned Tisserat, 2007

g y pjuglandis

• Killing black walnuts in CO, ID,OR, and UT

• Previously unknown Geosmithiasp. association in N. America

??

sp. association in N. America

• Black walnut comprises ~1.9% ofhardwoods available in the US

Oak WiltCeratocystis fagacearum

• Farthest east case found nearSchenectady, NY 2008

Bruce Moltzan 2008

y

• Lethal wilt disease of the red oak sub‐genus

• Oak wilt fungus vectored by sapfeeding beetles and root grafts

• Suppression programs using rootgraft disruption and systemicinjection are effective, but laborious

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Also, Abiotic Stresses are compounding and accelerating insect and pathogen attacks

Abiotic stresses such as

O• Ozone

• Acid Rain

• Heavy metals

• Drought

• Climate change

Nicholas, et al. 2000.

Many forest species have already been devastated by diseases or insects, and the list is growing.

58 million acres at risk to insect and disease.25% mortality in next 15 years (3x normal).y y ( )

Best known

• Loss of the American Chestnut from the blight.

• Devastation of American Elm by Dutch Elm disease.

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American Chestnut Foundation/Forest History Society

Chestnuts

Food for wildlife

Agriculture &Forestry

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Chestnut wood

R t i t tRot resistant

Easy to work

91,760,000 chestnut shingles made in 1909

Spread of the blight(Cryphonectria parasitica)

• The blight was discovered in 1904 in NY.

• Within 50 years it had spread throughout chestnut’s entire range killing 4 billion trees.

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Chestnut Blight

• Cryphonectria parasitica (fungus)

• Colonizes wounds

• Forms a canker & girdles tree

The American Chestnut Foundation

www.acf.org

Founded in 1983 as a non‐profit organization, largely supported by membership contributions.

Goal: To restore the American chestnut tree to the eastern U S h h b di d i hU.S. through breeding and cooperative research.

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TACF’s Backcross Breeding Program

Chinese

F1 ½ American (50%)

AmericanX

AmericanXEach generation select for: Blight resistance American characteristics

Final Product: Good resistance + American type tree

BC1 ¾ American (75%) American

BC2 7/8 American (88%) American

BC315/16 American (94%)

X

X

BC3X

American characteristics

3 /16 ( ) 3

BC3 F2 15/16 American (94%)

X

BC3 F2

BC3 F3 15/16 American (94%)

X

Genomic Tool Development For The Fagaceae

Principal Investigator: Ronald Sederoff1

Project Manager: Nick Wheeler1

Co‐PIs: Sandra Anagnostakis6 , John Carlson4 , Fred Hebard3 , Tom Kubisiak7, D hli Ni l 1 Willi P ll5 P l Si 3 Ch i S ith1 J ff T ki 2Dahlia Nielsen1 , William Powell5 , Paul Sisco3 , Chris Smith1 , Jeff Tomkins2

1 North Carolina State University, Raleigh, NC

2 Clemson University, Clemson, SC

3 The American Chestnut, Foundation, Asheville, NC

4 Pennsylvania State University, University Park, PA

5 State University of New York, Syracuse, NY

6 Connecticut Agricultural Experiment Station, New Haven, CT

7 SIFG‐USDA‐Forest Service, Saucier, MS

Genomic Tools for the FagaceaeProposal Development Meeting

March 23, 2004

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Goal: Develop resources for genomics and genetics for Fagaceae trees species, focusing on EST databases and DNA markers.


Species: Chinese chestnut: (Castanea mollissima)American chestnut (Castanea dentata)Northern red oak (Quercus rubra)White oak (Quercus alba)American beech (Fagus grandifolia)

Bead Based Amplification & P iPyrosequencing

• Read lengths app 350 bases

• > 400,000 reads per run

• Single‐read accuracy of > 99.5%

• Unigene accuracy of > 99.99%

M. Margulies et al., Nature 437, 376 (2005).

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EST databases and DNA markers for Chinese chestnut: (Castanea mollissima)American chestnut (Castanea dentata)Northern red oak (Quercus rubra)


Northern red oak (Quercus rubra)White oak (Quercus alba)American beech (Fagus grandifolia)

Genetic and Physical maps for Chinese chestnut

cDNA Libraries Sequenced:(1) Infected stem tissue for Chinese chestnut cv. Nanking (TACF) (GS20)

(2) Infected stem tissue for American chestnut cv BA69 (TACF) (GS20)

(3) Uninfected stem tissue from Chinese chestnut cv Mahogany (TACF)

(4) Uninfected stem tissue from Chinese chestnut cv Nanking (TACF)

(5) Uninfected stem tissue from American chestnut Wisneiwski (CAES)

(6) Uninfected stem tissue from American chestnut Watertown (CAES)

( ) h l l f h h k ( )(7) Whole plant tissues for Chinese chestnut Nanking GR119 (TACF)

(8) Whole plant tissues for two Chinese chestnut Mahogany BX316 (TACF)

(9) Whole plant tissue for American chestnut Wisneiwski (CAES)

(10) Whole plant tissue for American chestnut Watertown (CAES)

(11) Northern red oak tree – Above ground tissues (PSU)

(12) Northern red oak seedlings –Roots (PSU)

(13) White oak tree ‐ Above ground tissues (PSU)

(14) White oak tree seedlings – Roots (PSU)

(15) Whole plant tissues for healthy American beech tree 1504 (USFS)( ) p y ( )

(16) Whole plant tissues for infected American beech tree 1506 (USFS)

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Species Mbp Contigs fl cds

Chi Ch 172 32 738 874

Transcript Contigs from the “Genomic Tool Development for the Fagaceae Project”

Chinese Chestnut 172 32,738 874

American Chestnut 146 46,364 344

Northern Red Oak 66.3 17,502 361

White Oak 48.0 13,626 311

American Beech 14.6 4,745 50

Total: 447 91 325 1679Total: 447. 91,325 1679

American Chestnut Canker Chinese Chestnut Canker

Comparison of American and Chinese Chestnut UnigenesGene Ontology Analysis for Biological Processes

Genes involved in response to environmental stimuli and stresses were more highly expressed in Chinese chestnut than in American chestnut

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Contig Name log2 noFunction

CCall_contig26701_v25.63 1-aminocyclopropane-1-carboxylate oxidase

CCall_contig35157_v25.20 MYB3R- and R2R3- type MYB- encoding genes

CCall_contig13503_v21.04 cytosolic ascorbate peroxidase APX1

CCall_contig28632_v23.07 allene oxide cyclase

CCall_contig27982_v22.90 ABC transporter family

CCall_contig42826_v21.43 cinnamoyl CoA reductase

CCall_contig13029_v21.78 JAZ1 is a nuclear-localized protein

Disease Response Genes Differentially Expressed in Chinese Chestnut Canker

CCall_contig9319_v2 1.12 subunit 6b of cytochrome c oxidase

CCall_contig8687_v2 0.72 isoform of 4-coumarate:CoA ligase (4CL)

CCall_contig38717_v26.44 lipoxygenase

CCall_contig3555_v2 4.90 MLP-LIKE PROTEIN 34 (MLP34)

CCall_contig24182_v21.84 12-oxophytodienoate reductase

CCall_contig43865_v22.00 cinnamate-4-hydroxylase

CCall_contig22956_v27.48 PR (pathogenesis-related) peptide

CCall_contig46205_v25.00 alpha-dioxygenase

CCall_contig8907_v2 3.28 similar to the antifungal chitin-binding

CCall_contig41513_v21.89 manganese superoxide dismutase (MSD1)

CCall_contig3523_v2 1.28 A senescence-associated gene

CC ll ti 39986 21 44 t i t i i hibit ti it H i lli

: Pathogenesis related

: Phenylpropanoid pathway

Barakat et al. (manuscript in preparation)

CCall_contig39986_v21.44 cysteine proteinase inhibitor activity

CCall_contig42078_v23.69 basic chitinase

CCall_contig46415_v24.78 dienelactone hydrolase family protein

CCall_contig5675_v2 0.90 glutathione peroxidase.

CCall_contig11269_v22.33 sinapic acid:UDP-glucose glucosyltransferase

CCall_contig32533_v21.16 MAP kinase kinase 2

CCall_contig38534_v23.66 flavonol 7-O-glucosyltransferase (EC 2.4.1.237)

CCall_contig4088_v2 4.52 ELICITOR-ACTIVATED GENE 3-1 (ELI3-1)

CCall_contig14020_v21.82 Transcription factor of the B-ZIP family

CCall_contig46027_v26.20 protein with ABA 8'-hydroxylase activity

: Hormone signalling

Chestnut candidate genes for resistance

• HydrolasesC tit ti d f• Constitutive defenses

• Jasmonic-acid dependent defenses• Signaling• Cell Wall biosynthesis genes• Hypersensitivity• Hypersensitivity• Systemic Resistance

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The chestnut genetic and physical maps from the Genomic Tool Development for the Fagaceae Project

~1100 SNPs + 250 SSRs

Tom Kubisiak , USFS

1100 SNPs + 250 SSRs750 cM, avg spacing 0.5 cM

Bert Abbott, CUGI

126,445 clones input, 18X genome depth1,377 contigs, 12,919 singletons1026 markers anchored

1) EST‐based SSRs: About 800 SSRs identified• Genetic Linkage mapping• Comparative genomics• Population structure studies• Integrating genetic and physical maps

Applications for chestnut genomics resources

Integrating genetic and physical maps

2) SNPs: About 12,000 SNPs identified• Discover by data pipeline, visual inspection, re‐sequencing• No need for sequence confirmation due to large numbers of reads• 3410 Species specific SNPs discovered in 1716 genes• Assayed by Illumina Golden Gate system for mapping and genetic

association studies

3) Candidate genes for blight resistance (202 being mapped)• Discover by Blast annotation• Validate by RT‐PCR• Test by transformation

4) Genome sequencing

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THE FOREST HEALTH INITIATIVE

Advancing Forest Health through Biotechnology

A collaborative effort to advance role of biotechnologies to address today’s pressing forest health challenges, especially stress resistance,

Eco‐genomics, BG 2011

y p g g , p y ,starting with restoration American chestnut.

http://www.foresthealthinitiative.org/

The Forest Health Initiative’s Biological Sciences projects:

Genomics• Whole genome sequence of blight‐resistant Chinese chestnut• Resequencing of American chestnut genome

Eco‐genomics, BG 2011

Genetic Technologies and Breeding

• Discovery of host candidate genes resistance

• Refine genetic maps and QTL for blight resistance• Map QTL for Phytophthora cinammomi resistance• Develop DNA markers to aid selection in breeding programs• Develop early blight screening protocol

Transgenics and Propagation• Refine somatic embryogenesis and transformation protocols• Test new candidate genes in transgenic plants• Screen transgenic plants in lab and field tests• Develop strategy for deploying transgenic trees in breeding

and ecological restoration programs

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Goals:

1) Develop a complete genome sequence for chestnut

The Chinese Chestnut Genome Sequencing Project

1) Develop a complete genome sequence for chestnut

2) Identify all genes in the three blight resistance QTL

3) Deliver candidate genes for blight resistance to

transgenic and genetic technologies projects

4) Provide the genome as a community resource

) h i l f i dd5) Demonstrate the potential of genomics to address forest health and ecosystem restoration.

Scientific Approach:

1. Develop a high quality reference genome sequence for Chinese chestnut (Castanea mollissima) cv Vanuxem by 454 technology


c est ut (Casta ea o ss a) c a u e by 5 tec o ogy

2. Assemble the genome into contigs and scaffolds

3. Assemble larger scaffolds using the genetic linkage map

4. Correct gene models and sequence gaps with Illumina data

5. Use the physical map to assemble pseudo chromosomes

6. “Resequence” additional chestnut genomes by Illumina

• Other Chinese chestnut genotypes for SNPs

• American chestnut for QTL loci comparisons

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Progress to date:

• 454 shot‐gun sequence at 18X* depth (14.2 Gb)

• Illumina sequence at 47X* depth (37 6 Gb)


Illumina sequence at 47X depth (37.6 Gb)

• 454 paired‐ends at 4X depth (3.6 Gb) from 3, 4‐7, 8, & 15‐20kb inserts

• Preliminary assemblies conducted at 1, 10, 15, 18X. Current build:‐ 454 sg and pe reads: 587,208,063 bp in 51,766 scaffolds‐ Illumina data at 47X: 620,212,200 bp in 1,372,438 scaffolds

• ‘Contamination’ check:

‐ 0.19% contigs and 0.35% reads align to mtDNA;

‐ 0.11% contigs and 0.28% reads mapped to chloro DNA.g pp

‐ 8.5% to 12.3 % of reads align to conserved repeat DNAs

• Transcript Alignments (EST unigenes)

‐ 96% of the NSF project Chinese chestnut transcript contigs

aligned with 98% identity to current genome assemblies

* depth shown above assumes a genome size for chestnut of app 800 Mbp

The 900+ Million base pairs of chestnut genome sequence can be pieced together using


can be pieced together using the genetic and physical maps

of each chromosomes.

Since we know where the three chestnut blight resistance loci are on the genetic map, we can use this alignment of maps and genome sequence to Identify all of h f bli h i

Scaffolds in burgundy; EST‐based loci from linkage map in red; Physical map contigs in blue.

the genes for blight resistance.

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How can Genomics and Bioinformatics help address current Forest Health Issues?

Challenges associated with forest trees:• Multiple stresses• Multiple stresses

• Multiple species in complex communities

• Large populations covering expansive areas

• Not a lot of time for traditional approaches

• Immense genetic diversity and adaptation

• Very low LD values y

• Few genetic tools available

Many species, how to chose?

• Phylogenetic breadth

• Ecological importance

• Economic importance

Angiosperms

Monocots

Magnoliales

Amborellales

Laurales

• Existing genetic resources

Provenance tests

Germplasm collections

Orchards and plantations

Limited or no genomics resources

• Acute biotic or abiotic threats

Eudicots

re eudiocts

Rosids

Proteales

FabalesRosalesFagalesMyrtalesBrassicales

l l

Malphigiales

Cor R

Asterids

Lamiales

Malvales

CornalesSapindales

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Chestnut LG_G vs peach LG 1 and 8

Comparative mapping with peach genome identified regions on peach G6 and G8responsible for powdery mildew resistance overlapped with corresponding chestnut QTL regions for blight resistance on chestnut LG_B and LG_G.

Foulongne et al., 2003.

Molecular Breeding 12:33‐50

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Best matches of proteins from the chestnut genome assembly are to peach and other related species

Only 1% of best matches to Arabidopsis.


The peach genome is best for chestnut gene discovery.

• peach, 23%• rice, 12%• grapevine, 7%• Eurosids 1 species, 56%

Best matches:

BLASTx alignments to model plant genomes in Phytozome

Comparative Genomics of Environmental Stress Responses in North American Hardwoods

Genomics and Genetics Resources to address Forest Health Issues

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Objectives:

Comparative Genomics of Environmental Stress Responses in North American Hardwoods

• Deep EST databases

• Reference populations for mapping and GWAS.

• Genetic linkage maps

• BAC libraries

• EST-based DNA markers for growth and stress-response traits.

• Identify syntenies among hardwood tree species• Identify syntenies among hardwood tree species.

• Efficient, inexpensive platforms for high throughput genotyping.

Plants we are studying• Liriodendron tulipifera

Model magnoliidFast growing hardwood

• Liquidambar styracifluaBasal EudicotFast growing hardwood

• Gleditsia triacanthos

Angiosperms

Monocots

Magnoliales

Amborellales

Laurales

• Gleditsia triacanthosWidely adapted nitrogen fixerPotential in agroforestry

• Quercus rubraDominant upland speciesComplementary to European workFine hardwood

• Juglans nigraCultivars/specialty cropAgroforestry programFine hardwood

Eudicots

ore eudiocts

Rosids

Proteales

FabalesRosalesFagalesMyrtalesBrassicalesMalvales

Malphigiales

Vitales

Fine hardwood

• Acer saccharumShade tolerantWidely adapted

•Fraxinus pennsylvannicaVery widely adaptedAsteridAcutely threatened

C

Asterids

Lamiales

Malvales

CornalesSapindales

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Liriodendron tulipifera

Acersaccharum

Native ranges of selected species

Liquidambarstyraciflua

Juglansnigra

Gleditsiatricanthos

Quercusrubra

Fraxinuspennsylvannica

Order SpeciesFramework

MapDense Maps

ESTsBAC

libraries

MagnolialesYellow-poplar Liriodendron tulipifera

X *

Sweetgum

Tool development objectives for each species

ProtealesSweetgum Liquidambar styraciflua

X X X

FabalesHoney locust Gleditsia triacanthos

X X X

Fagales

Black Walnut Juglans nigra

X X X

Northern red oak Quercus rubra

X X X

SapindalesSugar maple Acer saccharum

X X

LamialesGreen ash Fraxinus pennsylvanica

X X

*, Resources available from previous NSF Plant Genome Research Projects

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EST Database Development ‐ Stress treatments of full‐sib seedlings for RNA

Treatment type

Treatment description

Treatment lengthtype description

Cold 4 °COvernight at 4 °C; 24 hours at normal temps

Heat 40 °C 24 hours

Drought To pre-dawn Drought water potential at 1.2 ~ -1.5 Mpa

App. one week

Wounding 4 holes per leaflet with paper punch

5 or 24 hrs

By “Progeny Exclusion” :

Identification of full sib mapping populations in

OUR APPROACH TO CREATING MAPPING POPULATIONS WITHOUT MAKING CROSSES

open‐pollinated seed by DNA marker analysis.

How many of the acorns on this tree are full sibs?

• Genotype all possible • Genotype all possible parent trees

• Genotype seedlings• Match seedlings to

parentsRomero‐Severson

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Jeanne’s Q. rubra mapping parents

Progeny Exclusion Approach:

We are identifying full sib mapping populations inWe are identifying full sib mapping populations in open‐pollinated seed by DNA marker analysis. We are

observing full‐sib frequencies up to 90%.

SM1:Female parentSM2: Male parentRomero‐Severson

Orchard locations for mapping populations

SpeciesSpecies LocationLocation PIPI

Quercus rubra Tennessee Scott Schlarbaum

Juglans nigra Missouri Mark Coggeshall

Liriodendron tulipifera Tennessee Scott Schlarbaum

Liquidambar styraciflua Missouri Mark Coggeshall

Gleditsia tricanthos Tennessee Scott Schlarbaum

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ngiosperms

Monocots

Magnoliales

Amborellales

LauralesMagnoliales: Yellow poplar(Liriodendron tulipifera)

Where are we now with population development?

An

Eudicots

e eu

diocts

sids

Proteales

FabalesRosalesFagalesMyrtales

Malphigiales

Vitales

Core

Ros

Asterids

Myrtales

Lamiales

BrassicalesMalvales

CornalesSapindales

ngiosperms

Monocots

Magnoliales

Amborellales

LauralesProteales: Sweetgum (Liquidambar sytraciflua)


An

Eudicots

e eu

diocts

sids

Proteales


Malphigiales

Vitales

Core

Ros

Asterids

Myrtales

Lamales

BrassicalesMalvales

CornalesSapindales

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ngiosperms

Monocots

Magnoliales

Amborellales

LauralesFabales: Honeylocust(Gleditsia tricanthos)


An

Eudicots

e eu

diocts

sids

Proteales


Malphigiales

Vitales

2009

Core

Ros

Asterids

Myrtales

Lamiales

BrassicalesMalvales

CornalesSapindales

2009

Established 2011East TN State Nursery

ngiosperms

Monocots

Magnoliales

Amborellales

LauralesFagales: Northern red oak (Quercus rubra)


An

Eudicots

e eu

diocts

sids

Proteales


Malphigiales

Vitales

Core

Ros

Asterids

Myrtales

Lamiales

BrassicalesMalvales

CornalesSapindales

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ngiosperms

Monocots

Magnoliales

Amborellales

LauralesFagales: Black walnut(Juglans nigra)

‘Schessler’ ‘Sparrow’


An

Eudicots

e eu

diocts

sids

Proteales


Malphigiales

Vitales

Core

Ros

Asterids

Myrtales

Lamiales

BrassicalesMalvales

CornalesSapindales

230 ‘Sparrow’ x ‘Schessler’ full sibs established to date

Coggeshall


giosperm

s

Monocots

Magnoliales

Amborellales

LauralesSapindales: Sugar maple(Acer saccharum)

Ang

Eudicot

se eu

diocts

sids

M

Proteales


Malphigiales

Vitales

Core

Ros

Asterids

Myrtales

Lamiales

BrassicalesMalvales

CornalesSapindales

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ngiosperms

Monocots

Magnoliales

Amborellales

LauralesLamiales: Green ash (Fraxinus pennsylvanica)


An

Eudicots

e eu

diocts

sids

Proteales


Malphigiales

Vitales

Core

Ros

Asterids

Myrtales

Lamiales

BrassicalesMalvales

CornalesSapindales

Coggeshall

Black Walnut BAC library prepared

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UNDERGRADUATE RESEARCH INTERNSHIP

AT PENN STATE , 2011

D t i G d Ch i t N l ith D K Sh k

OUTREACH, EDUCATION

Dantria Grace and Christen Nelms, with Dr. K Shumaker

Hardwood Genomics Project Newsletters

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www.hardwoodgenomics.org

Ecological and evolutionary genomics research can identify the genes responsible for variation in adaptive traits in natural populations, including stress resistance.

This will provide new opportunities to address forestThis will provide new opportunities to address forest health threats, starting with GWAS for environmental stress resistance.

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Association Genetics can illuminate the basis of adaptive gene complexes

in natural populations

• What classes of genes (regulatory or structural) provide most of the

adaptive variation?

• Are the same loci involved at different evolutionary levels (within‐

population, among‐population, and among species)?

• How does natural selection affect molecular diversity and linkageHow does natural selection affect molecular diversity and linkage

disequilibrium?

The Chestnut Genome Project

Penn StateStephan Schuster, Webb Miller

Richard Burhans, Charles Addo-QuayeJi Qi, Lynn Tomsho, Lindsay Kasson,

Tyler Wagner, Nicole ZembowerClemson University

Penn StateCharles Addo-Quaye, Teo Best,Tyler Wagner, Nicole Zembower

Co-PIs

Jeanne Romero-Severson, University of Notre Dame

Scott Schlarbaum, University Tennessee at Knoxville

Hardwoods Genomics Project

Acknowledgements

yBert Abbott, Steve Ficklin Meg Staton, Chris Saski

Scott Schlarbaum, University Tennessee at Knoxville

Mark Coggeshall, University of Missouri

Haiying Liang, Clemson University

Oliver Gailing, Michigan Technological University

Ketia Shumaker, University of West Alabama

Meg Staton, Clemson University

Nick Wheeler, Project manager

Genomic Tool Development for the Fagaceae Project

P St tPenn StateJohn CarlsonAli Barakatm Haiying Liang, Ji Qi,Stephan Schuster, Lynn Tomsho, Scott DiLoreto, Tyler Wagner, Kerr Wall

SUNY ESF, Syracuse:William Powell, Kathleen Baier

The American Chestnut Foundation:Fred Hebard , Paul Sisco

CLEMSON Genomics Institute Bert Abbott, Barbara Blackmun, Chun-HuaiCheng, Eric Guang-Chen Fang, Steve Ficklin, Meg Staton, Jeff Tomkins

NC State UniversityRon Sederoff, Nick WheelerChris Smith. Dahlia NielsenConnecticut Agricultural Expt. StationSandy Anagnostakis, Lila Pinchot

USDA/ Forest ServiceTom Kubisiak, Dana Nelson

Genomics of Environmental Stress Responses in Hardwoods€¦ · 10/5/2011 6 1000 Cankers Disease...

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