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BY RICHARD A. SNIEZKO AND JIM SMITH Trees in our native and urban forests and forest plantations are invaluable for our survival and well- being. Understanding and utilizing the nat- ural genetic varia- tion that exists in our tree species allows us to manage some of the impacts from the biotic and abiotic stresses that trees face and to guide our future reforestation and restoration efforts. The October 1995 issue of the Western Forester focused on forest genetics and its application in manage- ment of our nation’s forests, particular- ly in the Pacific Northwest (PNW). This vital work has continued in the ensuing 22 years, and in this 2017 issue some of those efforts are presented and expanded to include some work in the Inland Empire. With several PNW groups recently celebrating 50 years (and even 100 years for early research) of forest genetic and tree improvement related activities (see articles by St. Clair, Erickson, Crawford, Yanchuk), it is timely to note accomplishments and continued needs. With a changing cli- mate and continuing accidental intro- ductions of destructive non-native insects and diseases, it is the natural genetic variability within our native tree species that will be the basic resource to help restore unhealthy forests, maximize productivity of plan- tation forests, and help meet the diverse future needs of society. Growing public awareness of genetics in general has brought forest genetics onto the list of forest resources that require active manage- ment and stewardship. Concepts of adaptation, innate disease or insect resistance (or susceptibility), and growth potential are now commonly understood to be largely controlled by genetic factors. Genetic conservation strategies are an important compo- nent of managing our food crop species, and they are equally impor- tant for our tree species. However, unlike most crop species, genetic con- siderations for some tree species involve not only their potential com- mercial use, but also the need for these species to continue to persist and evolve for generations in natural ecosystems. The awareness of forest genetics and value of tree improve- ment has been embraced by federal, state, county, tribal, and private land managers. As an investment, applied forest Back to the Future: Genetics and our Northwest Forests SOCIETY OF AMERICAN FORESTERS (CONTINUED ON PAGE 2) September/October 2017 Oregon • Washington State • Inland Empire • Alaska Societies Volume 62 • Number 3 Western Forester In This Issue: 50 Years of Forest Genetics in the Northwest PHOTO COURTESY OF JIM SMITH Thirty-year-old Douglas-fir progeny test 10 years after thinning on Drift Creek near Toledo, Ore. Fertilizer and thinning effects were studied along with the impact of harvest machine travel. From this and subsequent tests at multiple sites, the best parent of the 47 tested here will produce 51% greater volume than average wild sources. Richard Sniezko Jim Smith

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BY RICHARD A. SNIEZKO ANDJIM SMITH

Trees in our nativeand urban forestsand forest plantationsare invaluable for oursurvival and well-being. Understandingand utilizing the nat-ural genetic varia-tion that exists inour tree speciesallows us to managesome of the impactsfrom the biotic andabiotic stresses thattrees face and toguide our futurereforestation andrestoration efforts.

The October 1995 issue of theWestern Forester focused on forestgenetics and its application in manage-ment of our nation’s forests, particular-ly in the Pacific Northwest (PNW). Thisvital work has continued in the ensuing22 years, and in this 2017 issue some ofthose efforts are presented andexpanded to include some work in theInland Empire. With several PNWgroups recently celebrating 50 years(and even 100 years for early research)of forest genetic and tree improvementrelated activities (see articles by St.Clair, Erickson, Crawford, Yanchuk), itis timely to note accomplishments andcontinued needs. With a changing cli-mate and continuing accidental intro-ductions of destructive non-nativeinsects and diseases, it is the natural

genetic variability within our nativetree species that will be the basicresource to help restore unhealthyforests, maximize productivity of plan-tation forests, and help meet thediverse future needs of society.

Growing public awareness ofgenetics in general has brought forestgenetics onto the list of forestresources that require active manage-ment and stewardship. Concepts ofadaptation, innate disease or insectresistance (or susceptibility), andgrowth potential are now commonly

understood to be largely controlled bygenetic factors. Genetic conservationstrategies are an important compo-nent of managing our food cropspecies, and they are equally impor-tant for our tree species. However,unlike most crop species, genetic con-siderations for some tree speciesinvolve not only their potential com-mercial use, but also the need for thesespecies to continue to persist andevolve for generations in naturalecosystems. The awareness of forestgenetics and value of tree improve-ment has been embraced by federal,state, county, tribal, and private landmanagers.

As an investment, applied forest

Back to the Future: Geneticsand our Northwest Forests

S O C I E T Y O F A M E R I C A N F O R E S T E R S

(CONTINUED ON PAGE 2)

September/October 2017 Oregon • Washington State • Inland Empire • Alaska Societies Volume 62 • Number 3

Western Forester

In This Issue: 50 Years of Forest Genetics in the Northwest

PHOTO COURTESY OF JIM SMITH

Thirty-year-old Douglas-fir progeny test 10 years after thinning on DriftCreek near Toledo, Ore. Fertilizer and thinning effects were studied alongwith the impact of harvest machine travel. From this and subsequent testsat multiple sites, the best parent of the 47 tested here will produce 51%greater volume than average wild sources.

Richard Sniezko

Jim Smith

tree breeding programs are unique inthat they are cumulative. Once parentsof interest are identified and incorpo-rated into seed orchards, gainsachieved in traits such as growth rate,stem form, or disease resistance will beavailable for the next forest plantationwithout annual input, unlike recurring

costs such as planting or fertilizing.Using tested parents as the basis forthe next generation of improvementprovides an ever-increasing catalog ofdata to help make future selections.What we do now will influence forestswell into the future.

The seed collected, the field trialsestablished, and the data gathered inforest genetic tests and tree improve-ment plantings provide baseline infor-mation on genetic variation in each of

our native tree species and which sitesthe different populations (provenancesor geographic sources) within a speciesmay be best adapted (Harrington,Howe). As part of applied forest genet-ics programs, seed has been collectedfrom tens of thousands of individualparent trees in the Pacific Northwestand Inland Empire, and many of theparent trees have been cloned by graft-ing into seed orchards or clone banks(Cress). Most of these selections arefrom the conifer species of highest eco-nomic importance, such as Douglas-firand ponderosa pine, but extensive col-lections of seed and vegetative materi-als (for grafting or rooted cuttings)have also been made for species suchas whitebark pine and Port-Orford-cedar. The seed of many of thesespecies can be stored for at least sever-al decades and provides a means of exsitu genetic conservation (a backupplan in case of disasters). The storedseed of individual trees provides animmediate source of material to evalu-ate genetic variation within thesespecies as new disease or insect threatsarise or as abiotic stresses from achanging climate become apparent.

While the level of federal fundingfor many applied tree improvementprograms has seen severe cuts sincethe 1980s, the contributions of bothgenetic materials and expertise devel-oped in those programs have contin-ued to be major factors in the region.

The large federal and state-ownedacreage in the regions is well repre-sented with selections of most majortree species, which often formed thenucleus of cooperative breeding pro-grams for their neighbors in the pri-vate sector (see various articles oncooperatives). While most agencieshave been limited in their ability toprovide financial support in recentyears, availability of resources and in-kind arrangements have been vital tothe success of many co-ops. In manycases, thanks to these programs, a reli-able supply of genetically diverse seedis available to meet the needs of eachbreeding zone for some of the com-mercial species. The federal agenciesgenerally have their own sources ofseed as do members of cooperatives,

2 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

Next Issue: Evolution of Land Ownership and Management Implications

Genetics and ourNorthwest Forests(CONTINUED FROM FRONT PAGE)

Western ForesterSociety of American Foresters

4033 S.W. Canyon Rd. • Portland, OR 97221 • 503-224-8046 • Fax 503-226-2515http://www.nwoffice.forestry.org/northwest-office/western-forester-archive

Editor: Lori Rasor, [email protected] Forester is published four times a year by the Oregon, Washington State,

Inland Empire, and Alaska Societies’ SAF Northwest Office

State Society Chairs

Oregon: Werner Krueger, 777 NW GardenValley Blvd., Roseburg, OR 97471;541-464-3277; [email protected]

Washington State: Wendy Sammarco,11707 346th Ave. NE, Carnation, WA, 98014;206-233-1569; [email protected]

Inland Empire: Bill Love, CF, Inland ForestManagement, 214 S. Center Valley Rd.,Sandpoint, ID 83864-9542; 208-263-9420;[email protected]

Alaska: Jeremy Douse CF, Tanana ChiefsConference, 122 1st Avenue, Ste. 600,Fairbanks, AK 99701; 907-452-8251 x3374;[email protected]

Northwest Board Members

District 1: Keith Blatner, Professor andProgram Leader for Forestry, School of theEnvironment, Washington State University,Pullman, WA 99164-99164; 509-595-0399 (c);509-335-4499 (o); [email protected]

District 2: Mike Cloughesy, Oregon ForestResources Institute, PO Box 463, Silverton,OR 97381; 503-329-1014; [email protected]

Please send change of address to:Society of American Foresters

10100 Laureate WayBethesda, MD 20814

[email protected]

Anyone is at liberty to make fair use of the material in this publication. To reprint or make multiple reproduc-tions, permission must be obtained from the editor. Proper notice of copyright and credit to the WesternForester must appear on all copies made. Permission is granted to quote from the Western Forester if thecustomary acknowledgement accompanies the quote.

Other than general editing, the articles appearing in this publication have not been peer reviewed for techni-cal accuracy. The individual authors are primarily responsible for the content and opinions expressed herein.

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but for other landowners, there areoptions such as Oregon Department ofForestry’s Seed Bank (see sidebar onpage 4).

Most of the early work in forestgenetics was undertaken with speciesof the highest commercial importance.However, understanding the geneticvariation in other tree species such aswhitebark pine, Port-Orford-cedar, fox-tail pine and Pacific madrone have alsobeen recently undertaken. The expert-ise and infrastructure developed for thecommercially important species hashelped make the work with thesespecies more efficient. In some cases,such as with whitebark pine, restora-tion plantings on federal lands alsoserve as genetic trials and as sentinelsto monitor the impacts of new bioticagents and abiotic events. Althoughcooperatives have been key organizersof tree improvement efforts for majorcommercial tree species(Jayawickrama, Rust), the USFS hasbeen the lead in programs to develop

genetic resistance to non-native dis-eases such as white pine blister rustand Port-Orford-cedar root diseaseresistance (Sniezko). But, even in thesecases, cooperators and partners havebeen key in the progress, and theimportance of sharing the result(resistant seed) is paramount to main-taining these species in our forests.Ongoing work in these resistance pro-grams will produce seed with evenhigher levels of resistance for restora-tion or reforestation.

Articles in the 1995 issue (posted atwww.nwoffice.forestry.org/northwest-office/western-forester/2017) coveredtopics of importance to forest geneticsand provides a synopsis of the earlyhistory, progress, and considerationsin many of the forest genetic and treeimprovement efforts needed to main-tain healthy and productive forests.

Forest genetic and tree improvementprograms are large and long-termundertakings (Lipow). Organizationshave banded together in the form of

cooperatives to carry out much of thiswork. In this issue, articles provideoverviews of tree improvement pro-grams, genetic resource management,and research efforts to provide thereader with knowledge of the history,benefits, and findings to help informfuture decisions.

By its nature, the future is oftenuncertain. However, the needs of societyfor products from our forests, and the

PHOTO COURTESY OF RICHARD SNIEZKO

Pacific madrone (Arbutus menziesii) provenance trial onStarker Forests land is assessed for growth, survival, andfoliage blight in fall 2015, four years after planting. A com-mon set of 105 families representing seven ecoregionsfrom central California to British Columbia were plantedon sites in California, Oregon, Washington, and BritishColumbia to evaluate the adaptive genetic variation in thishardwood species and serve as sentinel plantings. Theproject has been an informal cooperative project ofseveral pathologists (WSU, ODF) and geneticists (USFSand British Columbia Ministry of Forests, Lands andNatural Resource Operations), with field sites providedby several organizations.

DATA SUMMARY FROM DORENA GENETIC RESOURCE CENTER

Shown on the y axis, variation in fourth-year height (cm)within and between Pacific madrone families from theseven ecoregions (x axis) represented in the provenancetrial at the Starker Forests site (1-7 represent Cascades,Central CA Foothills and Coastal Mountains, Coast Range,Klamath Mountains, Puget Lowland, Sierra Nevada, andWillamette Valley ecoregions, respectively). Note the largevariation in height among families within the same eco-region (family = seedlings originating from seed of onemother tree).

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 3

Special Thanks

The Western Forester thanksGeorge McFadden of the Bureau ofLand Management for the idea topublish an issue focusing on 50years of forest genetics in theNorthwest.

George, BLM’s Oregon/Washington silviculturist, spear-headed the theme idea, coordinat-ed authors and topics, and provid-ed financial support to publishthis expanded 32-page issue.

The efforts of George and all theauthors is another example of thecooperative spirit in the forestryand scientific community. ◆

(CONTINUED ON NEXT PAGE)

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need for healthy forests for recreationand other services will continue to grow.Experience has taught us that we willlikely see new destructive non-nativepathogens and insects in our forests. Achanging climate will place stress onour forests. Knowledge of the geneticvariation of our tree species helps us tomeet the increasing future needs ofsociety and reduce the impacts of biotic

and abiotic stresses that will come. Forthe most part, past tree improvementefforts in western North America utilizeclassical selective breeding techniquesto increase genetic gain in traits of inter-est, but in the future a detailed under-standing of genomic resources will helpprovide more precise knowledge toguide breeding efforts (Howe). Trees areon the landscape for decades and even

hundreds of years or more. Forestersand those working to maintain foresthealth need to be thinking well into thefuture to ensure forests have the bestchance to meet ecological and societalneeds. Most of our planted forests havebeen influenced positively by the stew-ardship and contributions of the forestgenetic community. The past record hasbeen encouraging, and with continuedsupport of the organizations involved,forest geneticists and tree breedersworking with colleagues in silviculture,ecology, forest health, and other disci-plines will ensure this continues. ◆

Richard A. Sniezko is a geneticist forthe US Forest Service Dorena GeneticResource Center in Cottage Grove, Ore.An SAF member, he can be reached at541-767-5716 or [email protected] Smith worked as a tree improve-ment forester and orchard managerfor Georgia-Pacific and Plum CreekTimberlands in Cottage Grove for 39years (retired in 2017). He can bereached at 541-554-6461 [email protected].

4 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

Seed Available through the Oregon Seed Bank

The Oregon Seed Bank has long-standing agreements that allow it to obtainseed from a variety of breeding programs in the PNW including Oregon StateUniversity, US Forest Service, BLM, and Oregon Department of Forestry pro-grams. These advanced genetic selections may be selected for disease resistance,enhanced adaptability, increased growth rates, enhanced timber qualities, or acombination of traits. In addition, the Seed Bank has statutory authority to pur-chase seed from any seed orchards established at the Oregon Department ofForestry Schroeder Seed Orchard complex. These arrangements provide familyforest landowners equal access to seed. The Seed Bank sells seed of theseadvanced genetic selections, along with natural seed collections, from a varietyspecies and seed zones to nurseries in Oregon. These nurseries then produceseedlings that are available for sale to family forestland owners and other cus-tomers. Additional information about the program along with listings of seedavailability is located at www.oregon.gov/ODF/Working/Pages/Seed.aspx. ◆

BY DAN CRESS

lot of time andmoney has been

devoted to improv-ing a whole range oftree species. Cattlebreeders use similartechniques toimprove their dairyherds, but is it practical to addressgenetics during everyday silviculture?In a word: Definitely.

We plant high-elevation sites withnoble fir instead of grand fir: that’sgenetic selection. Douglas-fir isn’tresistant to laminated root rot, yetwhite pines and cedars are. Maybeplant the pines if you have access torust-resistant stock and if the rust pres-sure isn’t too severe. Maybe use cedardue to its higher value, especially ifthose materials have been selected forresistance to deer browse. Consciouslyor not, silviculturists have been utilizinggenetics for ages. Today’s technologyjust allows us to be even more targetedthan we have been in the past.

Tree improvement isn’t cheap. Analternative would be to avoid plantingentirely and hope for the best with nat-ural regen. There are places where thismight make sense, but this usually isn’tthe case here in the Pacific Northwest.Seed zones are available for guidingwoodsrun cone collections, so why dowe go to great lengths to actually growthe seed to grow the seedlings? Millionsof dollars are spent every year to createand manage seed orchards. We even goso far as to control the pollen that polli-nates each orchard cone. Orchards cangenerate large seed with vigorousembryos, big cotyledons, lots ofendosperm, and so on. Tree improve-ment programs are what we use toactually decide what is, or isn’t, includ-ed within these orchards. Think oforchards as an investment rather thanan expense; we wouldn’t use them ifthey didn’t make financial sense.

The biggest costs in tree improve-ment come from breeding and proge-ny testing. We hand-pollinate femaleflowers with pure pollen so we knowthe pedigree of every seed that we

sow to gen-erate thetestseedlings.Thousandsof them areoutplantedat a varietyof siteswhere theirperform-ance istracked fora decade ormore. Thevery bestselectionsfrom thosetests arewhat we useto establish newer and better orchardsover time.

Consider western hemlock from outon the coastal strip. The first-generationtests involved about 25,000 trees fromeach of five separate zones. The best ofthose materials went into seed orchardsand were also used as the parents in asecond-generation breeding program.Those pedigreed seedlings weredeployed at new test sites spread all theway from Newport to the northern tipof Vancouver Island. This means 83,000more selection options that can be usedto establish the next round of seedorchards. Want to select for growth atvarious locations and ages? Want toconsider stem form and wood proper-ties? Do Tillamook families growing atForks give us insights about how treesmight respond to climate change? Whatcrosses are we making for the third gen-eration and beyond? The math getscomplex, yet this is exactly the type ofwork that we do, for many species, on adaily basis.

What about seed zones? They implythat local seed will be the best seed. Acumbersome yet better descriptionwould be to say that “local seed is asafe bet in the absence of pedigreeddata.” Woodsrun seed collected from asite is presumably well-adapted to thatsite, but there could certainly be some-thing just as suited to that environ-ment that grows faster, has betterwood properties, has better stem form,

and so on. Tree improvement is whatwe use to compare all of these options.

We have sound data on the per-formance of each test tree and that ofits parents, its cousins, its siblings, etc.Great, so how does growth and adap-tation within a test site compare toperformance under operational condi-tions? Realized gain trials have beenestablished to address such questionsover time. By no great surprise, treeimprovement is a success here in thewest just like it has been for things likeradiata pine in New Zealand andloblolly pine in the south.

The next step is to adjust the variousgrowth and yield models so they taketree improvement into account.Another line of study is to optimize thetrade-offs between traits such as growth,stem form, wood properties, andgrowth rhythm. These past 50 years’worth of breeding and testing generated128,000 second- and third-generationselection candidates for some new seedorchards that I designed earlier thisyear. We can make substantial gains, foran array of traits, when working withpopulations this huge. I can only imag-ine what folks will have available tothem 50 years from now! ◆

Dan Cress, an SAF member, is a forestgeneticist, Regenetics Forest GeneticsConsulting, in Seattle, Wash. He can bereached at 206-310-8963 or [email protected].

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 5

Why Grow the Seed to Grow the Seedling?

A

PHOTO COURTESY OF DAN CRESS

An excellent Douglas-fir cone crop. The cones on these threebranches can produce about 10,000 seed.

BY KEITH JS JAYAWICKRAMAAND TERRANCE Z YE

he PacificNorthwest

(PNW), similar toother forestry areasgrowing temperateconifers, has a longhistory of appliedcooperative treeimprovement. Withmultiple landown-ers typically manag-ing forestland withina given geographicarea, it has been logi-cal to pool resourcesfor the expensive,long-term endeavorof field testing andbreeding. Treeimprovement leadership was first pro-vided in the PNW via the IndustrialForestry Association and the US ForestService PNW Research Station (jointlycoordinating the Progressive TreeImprovement System) from 1966 to1985 and after 1986 by the NorthwestTree Improvement Cooperative(NWTIC).

Establishment of first-cycle Douglas-fir tests began in 1967 and continued to1993. A typical first-cycle cooperativetested 200-300 trees selected at anintensity of about three trees per 1,000acres on 6-10 test sites. Local adapta-tion was emphasized leading to smallbreeding zones. In addition to testingprograms designed as cooperative pro-grams, four independent programs

(Bureau of Land Manage-ment, Georgia-Pacific,Simpson, Washington StateDepartment of NaturalResources) came later underthe NWTIC umbrella. About1,000 first-cycle Douglas-firtests were established, inwhich approximately 30,000first-generation selectionswere tested. The earliest full-sib second cycle test wasplanted in 1984, but the restwere planted after 1996 withestablishment scheduled tocontinue through 2022.Second-cycle Douglas-fir siteshave been established fromSkagit County in Washingtonnear the Canadian bordersouth to Curry and Douglascounties in Oregon (andeventually to the northwesttip of California).

Tree height was assessedfrom the first measurementsaround 1975, and diameter atbreast height (DBH) from thefirst age-10 measurements.Stem defect (forking, rami-corns, and stem sinuosity)and wood density were rou-tinely assessed from the early1990s after those traits wereshown to be heritable andmeasurement protocols established.Three additional traits (spring bud-break, second flushing, and fall coldhardiness) are now being scored insecond-cycle tests. Needle retention isassessed in the area of the northwestOregon coast exposed to Swiss needlecast disease. Non-destructive acoustictools for assessing wood stiffness intree improvement became availablearound 2005, and so far about 32,000trees have been assessed. Other traits(e.g., drought damage, top break) areassessed in certain tests.

Third-cycle breeding orchards wereestablished starting in 2006. All thesecond-cycle cooperatives decided toproceed to third-cycle breeding andtesting. Compared to a delay of 20-34years from the start of individual first-

cycle programs to establishment ofsecond-cycle tests, the goal is to estab-lish third-cycle tests within 15-20 yearsafter second-cycle tests were planted.The first group of third-cycle tests wasestablished in March 2017 on six siteson the Oregon coast.

Breeding and testing of westernhemlock has been essentially a small-scale version of the Douglas-fir effort.First-cycle testing began in 1975, and asecond-cycle program for the coastalstrip of Oregon, Washington, andsouthwest British Columbia started in1992. Age-10 data collection for allthese second-cycle sites, testing 539full-sib crosses, was completed by2009-10. Third-cycle breeding isunderway. There was also a small first-cycle testing program for noble fir.

6 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

Development of Cooperative Tree ImprovementPrograms in the Pacific Northwest

T

Keith JSJayawickrama

Terrance Z Ye

PHOTO COURTESY OF CASCADE TIMBER CONSULTING

Genetic gain demonstration from a second-cycle test showing row plantings of a goodcross, a cross near the population mean, andwoodsrun seedlings. Both height poles are setat the same height.

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Cooperative seed orchards play avital role in advancing tree improve-ment. Many cooperators have toosmall an annual seed need to justifyestablishing their own seed orchard;other larger entities do not wish to adda specialized task such as seed orchardmanagement, plus specialized staff,facilities, and equipment, to theirforestry operation. Since seed produc-tion takes 8-15 years and can be affect-ed by many factors (rainfall, soil type,incidence of frost, precipitation), it isoften far safer to co-locate an orchardblock on a known successful operationthan take chances on an unprovenpiece of ground. Economies of scalealso make concentrating seed produc-tion on a few productive sites moreefficient. The Oregon Department ofForestry and the Bureau of LandManagement have been the mainhosts of cooperative seed orchards.

Based on responses from an NWTICsurvey, the number of forest treeseedlings planted in western Oregonand Washington originating from treeimprovement programs in 2017(coastal DF, hemlock, ponderosa pine,noble fir, and western redcedar) stoodat about 65.2 million with about 10.0million woodsrun trees planted. The2017 species summary (in millions) was61.7 Douglas-fir, 7.9 western hemlock,2.1 noble fir, 2.4 ponderosa pine, and1.2 western redcedar.

NWTIC has a strong interest in real-istic rotation-age estimates of realizedgain in operational conditions, and as aresult are investing heavily in establish-ing gain trials. Age-20 data have beenobtained from the oldest realizedgenetic gain trial (Molalla, planted1997), with the elite treatment showing19.6% superiority over woodsrun involume per acre at a tighter spacing. Inthe younger trial (Grays Harbor, plant-ed 2005 and 2006), the best performingfull-sib cross had age-9 realized gains of19.9%, 20.3%, and 62.3%, respectively,for height, DBH and volume, based on480 progeny planted on six sites.Second-cycle realized gain trials arecurrently being established forDouglas-fir and western hemlock.

NWTIC is an umbrella unit provid-ing specialized, highly technical serv-ices that would be expensive and inef-ficient for each cooperator or regionalcooperative to provide on their own.

Data analysis is the most vital of thoseservices, and includes advanced BestLinear Unbiased Prediction analyses,which has made it possible to providepredicted genetic gains and combinedata across breeding zones or acrossgenerations. Detailed reports are pro-vided that explain, visualize, and inter-pret the results.

Computer simulation and data re-sampling are routinely conducted forseeking optimal breeding strategies.Management of tree improvement datais also important: a very large numberof records (including information on3.38 million first-generation and 0.67million second-cycle progeny) are nowhoused in an SQL Server database witha Microsoft Access interface.

Since 2001, NWTIC has aided theestablishment/upgrading of high-gain1.5-generation or second-generationproduction orchards, and has provid-ed candidate selection lists for most ofthem. This impacts a substantialamount of current and future reforesta-tion in western Oregon, Washington,and California. NWTIC faculty havebeen first authors or co-authors on 22peer-reviewed publications and 40 con-ference and meeting presentations orother reports since 2000. NWTIC hasalso organized or helped organizeworkshops and short courses to takerelevant, practical, technical informa-tion and make it accessible to busypractitioners in tree improvement andforestry. Finally, the NWTIC routinely

pools resources with other university-based cooperatives to address ques-tions important to plantation forestryin the region. For example, NWTIC, incollaboration with the PacificNorthwest Tree Improvement ResearchCooperative, has initiated a genomicselection project on Douglas-fir; thepreliminary results are promising.

Looking to the future, the nextdecade will see emphasis on establish-ing third-cycle progeny sites. We cananticipate some increase in speciesdiversification, with recent interest innoble fir and western redcedar.Cooperatives will need to adapt torapid changes in land ownership,which are generally disruptive to long-term projects; we look to find new par-ticipants for cooperative work—espe-cially stable landowners with a strongstake in long-term forest health, pro-ductivity, and value improvement. Tothe extent supported by data, it will bepossible to consolidate breeding zonesand make them more efficient, increas-ing gain per unit cost and time. ◆

Keith JS Jayawickrama is director of theNWTIC at the College of Forestry,Oregon State University, Corvallis,while Terrance Z Ye is quantitativegeneticist for the NWTIC. Keith can bereached at 541-737-8432 [email protected] can be reached at 541-737-9881 or [email protected].

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 7

BY MARC L. RUST

he InlandEmpire Tree

ImprovementCooperative (IETIC)was established in1968 by a group offoresters who recog-nized the need forgenetically improved seed sources toensure healthy and productive forestsfor the future. IETIC uses classicalplant breeding techniques includingselection, testing, and breeding toidentify superior genotypes for inclu-sion in production seed orchards.

Initial efforts were focused on pon-derosa pine (Pinus ponderosa var. pon-derosa) tree improvement and our firstprogeny tests were established in 1974.During that same year, members votedto expand IETIC to include additionalspecies (western white pine, westernlarch, Douglas-fir, and lodgepole pine).

During the early years, membersselected phenotypically superior“plus-trees,” collected open pollinatedseed for progeny testing, and estab-lished and measured progeny tests.Our first-generation progeny testshave provided valuable data andgenetic material for establishing seedand breeding orchards.

Recent efforts have focused on pro-viding opportunities for IETIC mem-bers to join forces to produce geneti-cally improved seed for their reforesta-tion programs. While a few of ourmembers had previously established

their own seed orchards with IETICgenetic materials, many of our mem-bers have relatively small ownershipsand the fixed costs associated withdeveloping their own seed orchardswere prohibitive. To remedy this,IETIC established new cost-share seedorchards that are funded by memberswho want to participate. Each mem-ber pays a percentage of the establish-ment and management costs andreceives an equivalent share of theseed produced in return.

The first cost-share seed orchardestablished was a western larchorchard designed to produce seedsuitable for eastern Washington andnorthern Idaho between 2,800 and4,200 feet in elevation. This orchardwas planted in 2007 and produced itsfirst sizable seed crop in 2014. Whilestill very young, the orchard has pro-duced seed each year for the pastthree years. Trees are topped to 2.5meters tall once they reach 4.0 metersin height. This technique promotes thedevelopment of pendent branches inlarch, which tend to produce moreflowers than typical lateral branches.

In addition, IETIC members haveestablished a cost-share ponderosapine orchard to produce seed for east-ern Washington and northern Idahofrom 2,400 to 3,800 feet elevation anda Douglas-fir orchard for 3,000 to 4,000feet elevation in the same region.Efforts are underway to identify suit-able sites for two more western larchcost-share orchards, one to serve high-er elevations in Washington and Idaho,

and one to serve member needs inMontana.

While these orchards are new, thecost share concept isn’t new to IETIC.For many years, IETIC membersshared costs to manage the R.T.Bingham western white pine seedorchard located in Moscow, Idaho.This orchard, originally developed as abreeding arboretum by the US ForestService, has produced more than 8,000pounds of blister rust resistant westernwhite pine seed for IETIC memberssince 1993. Seed from this orchard hasbeen deployed by members inWashington, Idaho, and Montana withgood success. Currently, efforts areunderway to improve the blister rustresistance level of the seed from thisorchard by: 1) establishing neworchard blocks using tested materialsfrom blister rust resistance screeningsconducted at the US Forest ServiceCoeur d’Alene Nursery; and 2) partici-pating with the US Forest Service insecond-generation breeding effortsdesigned to produce improved levelsof blister rust resistance for the future.

With our first generation of progenytesting largely completed and the seedorchards developed from them begin-ning to supply genetically improvedseed to our members, IETIC has begunto plan for the future. To ensure thatour best genotypes from the progeny

8 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

The IETIC: Five Decades of Tree Improvement

T

PHOTO COURTESY OF MARC L. RUST, IETIC

Western larch seed from the IETICcost-share seed orchard shortlyafter germination. Germinates willbe thinned to one per cell.

tests are always available, we havebeen grafting them onto rootstock andplanting them at the Russell H.Hudson Gene Archive located on theUniversity of Idaho ExperimentalForest. Russ was instrumental in theformation of IETIC and served as IETICchair for more than 20 years. His lead-ership and commitment to the pro-gram have had a lasting impression.

In addition to preserving our bestgenetic materials for the future, mem-bers have recently become interestedin launching new advanced generationbreeding programs for two of our mostimportant species, western larch andDouglas-fir. Breeding plans and oper-ating agreements have been draftedand are being reviewed by members.These plans, focused on a traditionalapproach of full-sib crossing to pro-duce offspring for progeny testing, arebeing designed to take advantage ofnew genomic-based tools and tech-niques as they become available.

As a small cooperative without adedicated research budget, IETIC hasestablished or funded several impor-tant studies to improve seed orchardmanagement techniques or developnew tools to help further our efforts todevelop improved seed sources for thefuture. Some highlights include:

1. Developing flower inductiontechniques to promote early floweringin western larch and ponderosa pineseed orchards. These techniques arebeing used in the Pullman PonderosaPine Seed Orchard with good success.

2. Establishing trials of stem inject-ed systemic insecticides aimed at con-trolling high-value seed crops in seedor breeding orchards that cannot beeasily treated using broadcast insecti-cide sprays.

3. Developing an illustrated publi-cation on the reproductive biology ofwestern larch to aid foresters and seedorchard workers interested in harvest-ing or producing larch seed crops. Thispublication is available at www.web-pages.uidaho.edu/ietic/.

4. Developing SNP (single nucleotidepolymorphic) markers for westernwhite pine in collaboration with col-leagues at Oregon State University.These genetic markers represent singlechanges in the base sequence at a par-ticular location on a chromosome. IfSNP markers can be found that are

highly correlated to the presence (orabsence) of particular blister rust resist-ance traits, they could potentiallystreamline tree breeding efforts by pro-viding early detection of genoptypeswith high (or low) levels of resistance.

5. Comparing growth characteris-tics of open pollinated clonal seed lotscollected from seven western larchclones in the IETIC seed orchard.While deployment of clonal seed lots iscommon in some species and regions,it has not been practiced to any signif-icant degree with native coniferspecies in our region. Seed was collect-ed in 2016 and recently sown for thisstudy. The study includes both a nurs-ery component to examine growth dif-ferences of seedlings and a long-termfield study where the clonal lots will beoutplanted in blocks along with anorchard bulk lot on three sites withtwo levels of competition control. Theresults of this study will help membersdecide if future crops from the orchardwill be collected and processed by

clone or bulked as an orchard lot.If there is one thing that is certain

about the future, it is uncertainty.Concerns about the impact of climatechange on native forests and the chal-lenges of competing in an increasinglylarge global marketplace will likelyintensify the importance of our treeimprovement efforts. A lot haschanged since IETIC was foundednearly five decades ago. However, ourmembers continue to realize theimportance of improved seed sourcesto ensure healthy and productiveforests for the future and are commit-ted to cooperative efforts to maintain abroad genetic base and achieve genet-ic gains through shared responsibili-ties and shared costs. ◆

Marc L. Rust is director of the InlandEmpire Tree Improvement Cooperativein Moscow, Idaho. He can be reached at208-885-7109 or [email protected].

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 9

PHOTOS COURTESY OF MARC L. RUST, IETIC

Left: The IETIC western larch cost-share seed orchard was established in2007. Ramets are topped to promote the development of pendant branchesand allow cones to be easily harvested using orchard ladders. Right: Maleand female flowers on a pendant branch.

FOREST RESOURCES TECHNOLOGY• SAF Accredited •

Ron Boldenow, Ph.D., C.F., ForestryRebecca Franklin, Ph.D., DendrochronologyBret Michalski, M.S., Wildlife Science

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10 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

BY ALVIN YANCHUK

rich history oftree improve-

ment in the PacificNorthwest exists,and BritishColumbia’s involve-ment started withthe pioneering workof Dr. Alan Orr-Ewing in the mid-1950swith inbreeding and wide-crossingstudies of coastal Douglas-fir. BritishColumbia’s Douglas-fir tree improve-ment program developed further inthe 1960s with an enormous con-trolled crossing program. Currently,coastal Douglas-fir seed orchards areproducing third-generation seedlingswith predicted volume gains of 25% at

rotation ages about 60. In the late 1960s and 1970s, British

Columbia’s work expanded to includeimprovement programs for interiorspruce and world-class provenancetesting—under the auspices of theInternational Union of Forest ResearchOrganizations—for lodgepole pine,true firs, Sitka spruce, coast and interi-or Douglas-fir, and other species oflesser commercial importance. Whileprovenance trials were done primarilyto identify superior provenances, thiswork was later instrumental for thedelineation of seed zones. During thisperiod, forest industry became muchmore interested in tree improvementand started to help the province withseed orchard establishment, test siteselection, and funding for research.Various cooperative “councils” wereformed to move the program forward,and for the past 20 years, the ForestGenetics Council of BC has served asthe over-arching coordinator, report-ing to the chief forester in BC, for treeimprovement (www.fgcouncil.bc.ca).

The heyday of tree improvement inBC, and actually, even around theworld, were the 1980s and 1990s.Universities were producing more for-est geneticists and there was a smallhiring pulse by the public and private

sector during that time. However, theBC government slowly changed itsfundamental funding structure forforestry investment: i.e., provincial for-est tree nurseries were privatized andfunding for tree improvement largelymoved to third-party “crown corpora-tion” agencies. Ironically, while wewere enduring more governmentdownsizing, there was pressure todevelop improvement programs forlodgepole pine, western larch, whitepine, and interior Douglas-fir as thenumber of trees being plantedincreased to over 200 million. Despitethese fiscal pressures, tree improve-ment programs progressed nicely, withfirst- and second-generation breedingand seed orchards now in place. In2017, over 260 million trees wereplanted in BC and about 75% of thesewere from seed orchards.

The science of tree improvementbecame more complex in the late1990s when issues like biologicaldiversity came to the forefront. Manyquestions were asked, including: “Howdo the products of tree improvementfit with provincial biodiversity conser-vation concerns?” and “How do weconserve genetic diversity in treeimprovement programs while breed-ing for gains in volume?” and “Howcan tree improvement programsmaintain or enhance adaptive capaci-ty to changing environments?” Toshow that species diversity was also apart of the genetic improvement pro-

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gram, projects were initiated for hard-woods (birch, alder, cottonwood,maple) and other softwood species ofminor commercial importance, suchas ponderosa pine. The establishmentof high-quality genetics field trials arethe mainstay of BC’s tree improvementprogram, and as of 2017, forest geneti-cists working with the BC governmenthave planted approximately 1,500 tri-als in the ground, ranging from 1 to 50years old, with 15 species.

Initially, early volume growth wasthe principal target trait for improve-ment in BC’s programs, but over timeother traits were included in theselection and testing programs. Forexample, wood density, although avery important trait for structuralwood properties, is now being meas-ured to ensure this key wood propertydoesn’t decline during selection andbreeding. During the 1990s manyspecies were tested for disease resist-ance and with some species, such aswhite pine, “silviculturally” usefulgains have been made for blister rustresistance. Over the last 20 years, treeimprovement programs have placedsignificantly more effort on screeningfor disease and pest resistance includ-ing screening for resistance to spruceweevil (e.g., Sitka and interior spruce),pine rusts, and deer browsing (westernredcedar), to mention a few. Theseefforts have focussed on using the cur-rent disease and pests we now face asa “venue” to look for more durableresistance features of trees that couldguard against future pests and dis-eases. New pests and diseases in east-ern North American forests, such asbirch bronze borer, butternut canker,and emerald ash borer, are examplesof the new forest health challenges wecan expect in the west. All in all, BCtree improvement programs are aimedat developing more durable trees!

Tree improvement programs in BCcontinue to be an important elementof managing and conserving the forestgenetics resource. Work is currentlyfocusing in three major areas:

1. Climate change. Our currentpopulations of trees may not be suit-able for future environments, and weare proposing to model future envi-ronments based on the suitable “cli-mate spaces” that each species hasbeen tracking over their history in BC.

The challenge here is to align the treescoming from our seed orchards to thenew projected climates zones in theprovince.

2. Genetic conservation. Whilebreeding is another form of geneticconservation (e.g., we are developingmultiple populations with multipletraits), geneticists are well aware of thevalue of the native gene pool thatforms the basis of forests in BC. Withclimate change, and the introductionof new pests or disease, gene conser-vation populations in reserves, clonebanks, or the seed center are valuableresources. It’s our form of forest insur-ance!

3. Innovative tree breeding science.Many new techniques are available tous, such as better analytical tools toevaluate the best parent trees, opti-mization software to balance geneticgain while minimizing inbreeding,more efficient field test designs,improved GIS modeling techniques forclimate predictions, LiDAR technolo-

gies for measuring trees in our trials,and better wood quality assessmenttools and genetic markers. Many of usenvy the next generation of tree breed-ers; these new tools will go a long wayto meet the challenges of climatechange and keeping forestry as a keyindustry in BC.

The future for tree improvementremains exciting. Despite the reduc-tion of forestry-related research in BC,the products of tree improvement arebeing widely used and appreciated,and most foresters view genetics as animportant tool to combat the chal-lenges that nature, and us humans,can throw at forests. ◆

Alvin Yanchuk works as a researchleader for Forest Genetics at theMinistry of Forests, Lands, and NaturalResource Operations in Victoria, BC. Hecan be reached at 250-387-3338 [email protected].

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 11

BY MICHAEL S. CRAWFORD

he Bureau ofLand Manage-

ment (BLM), anagency within theUnited StatesDepartment of theInterior, administers258 million acres ofpublic lands, mostly in Alaska andeleven western states. Of this landbase, 2.2 million acres in westernOregon are managed for forestry. Aspart of their effort to develop healthy,

fast growing sustainable forests, theBLM made a major commitment over50 years ago to tree improvement andforest genetics, understanding thelong-term commitment required tomeet their goals.

The BLM instigated tree improve-ment in 1965, establishing the HorningSeed Orchard on 320 acres south ofPortland, Ore. Initially operated as astand-alone program, it made selec-tions based only on phenotypic charac-teristics, which did not employ progenytesting to analyze tree qualities. In 1968the agency changed direction and

joined the Cooperative Progressive TreeImprovement Program (which mor-phed into the Northwest Tree Improve-ment Cooperative, or NWTIC) to widenthe genetic base for testing and helpshare the cost of progeny evaluationswith industrial forest partners. Theagency was highly involved in regionalbreeding programs and by 1987 hadinstalled 209 progeny test sites in west-ern Oregon. The BLM participated intree improvement programs toenhance growth-and-yield in Douglas-fir, western hemlock, and noble fir, aswell as working with the US ForestService Dorena Genetic ResourceCenter to promote resistance testing ofwhite pine blister rust in five-needlepines.

By the mid-1980s, the BLM greatlyexpanded their orchard footprint.Horning was expanded to cover theagency’s Douglas-fir, noble fir, westernhemlock, western white pine, westernredcedar, and sugar pine seed needs ofnorthwest Oregon. Two additionalDouglas-fir seed orchards were soondeveloped: Tyrrell covered the seedneeds of west-central Oregon and thesouthern coast, while Provolt grewseed for southwest Oregon. By theearly 1990s, the BLM managed anextraordinary 490 acres of Douglas-firseed production orchards in westernOregon to cover reforestation needs of1,300 lb/year.

Just as the BLM orchards were com-ing into production, a major change infederal forest management occurred,resulting in a dramatic decrease inseed needs for the agency. By the mid-1990s, it became apparent that thefirst-generation orchards, designed tomeet the seed needs for the harvestlevels of the 1970s and early 1980s,provided significantly more seed pro-duction potential than the BLM need-ed. As a result, the BLM developedtheir first seed orchard memorandumof understanding (MOU) at Tyrrell in1998, providing industrial forestlandowners and forest nurseries in westernOregon the opportunity to cooperatewith the BLM in improved seed pro-duction. Over the next four years, sim-ilar MOUs were in place for the otherthree BLM orchards.

12 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

BLM Genetics Program Prepares for the Future

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Development of BLM Seed Orchards

By the mid-1980s, the BLMhad estab-lished threeDouglas-firseed orchardfacilities andone sugar pineorchard inwesternOregon, man-aging nearly450 acres ofimprovedDouglas-firorchard blocks.These orchardscovered a 1,200lb/year seed need for the Salem, Eugene, Roseburg, Coos Bay, and MedfordBLM Districts. Six greenhouses at Horning and Sprague orchards producedabout 2.5 million containerized seedlings per year. ◆

PHOTO COURTESY OF BLM ARCHIVES

In 2006, decision makers recom-mended rightsizing the entire orchardprogram by closing Sprague andProvolt seed orchards in southwestOregon, downsizing orchard staff, andredesigning the tree improvement pro-gram to maintain quality while contin-uing to look for efficiencies. Horningand Tyrrell were to become the futurebase of operations for the program,reestablishing new orchard blockswith genetic material from the closedfacilities and developing advancedgeneration orchards as tested materialbecame available. First-generation

Douglas-fir breeding units in westernOregon were consolidated into morelogical geographic regions based onprogeny test analysis by NWTIC. Thisresulted in a significant reduction inthe number of acres required forDouglas-fir seed production, fallingfrom 460 acres in 1990 to 47 acres in2010. A combination of 14 elite 1.5-generation and 2.0-generationorchards replaced 45 first-generationDouglas-fir orchard blocks planted inthe 1980s.

Currently, the BLM has 26 coopera-tors in its seed production program

covering federal, state, industrial, andtribal forestland between northernCalifornia and western Washington.Cooperators provide proportionatefunding for their level of participationto cover all facets of seed orchardmanagement, resulting in significantyearly support from the private sectorto operate the facilities. In return, thecooperators receive their share ofimproved orchard seed. At Tyrrellalone, cooperators have received over11,000 pounds of improved Douglas-fir seed over the past 19 years.

With 50 years behind it, the BLMcontinues to prepare for tree improve-ment in the next half-century. Majorimprovements to the orchard infra-structure, including updates of offices,storage buildings, and water systems,were recently completed. A new green-house/shade house was constructedfor small-lot growouts and as a region-al grafting center for cooperators, anda newly completed 50-person confer-ence center will provide space forregional forestry and tree improve-ment meetings. An agency with a longhistory in working to improve forestgenetics, the BLM continues to staypresent and remain an active player intree improvement in the PacificNorthwest. ◆

Michael S. Crawford is BLM SeedOrchard Program manager, TyrrellSeed Orchard in Lorane, Ore. He canbe reached at 541-767-0460 [email protected].

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 13

BLM Orchards for the Future

Approximately 83 acres of newlyplanted advanced generationorchards have been developed since2010, which will help address thefuture seed needs of the BLM and its26 orchard cooperators. Speciesinclude Douglas-fir, noble fir, sugarpine, and western hemlock. As newgenetic material becomes availablefollowing analysis of breeding coop-erative progeny testing, the BLMwill establish updated orchards.Trees are paper mulched for earlyvegetation control and tubed to pro-tect grafted seedlings from deer andelk damage. Tree tags include bar-codes to assist with efficient inven-tory. The taller trees in the back-ground are part of a Port-Orford-cedar preservation orchard man-aged in conjunction with theDorena Genetic Resource Center,US Forest Service. ◆ PHOTO COURTESY OF M. CRAWFORD

BY VICKY ERICKSON

he PacificNorthwest

Region (Region 6) ofthe US Forest Serviceencompasses 26.2million acres inOregon andWashington. The 16national forests and grasslands supportsome of the most diverse ecosystemsand flora in the US, with habitats rang-ing from the high deserts east of theCascade Mountains in central Oregonto the rain forests of the OlympicPeninsula in western Washington.

The region’s genetics program waslaunched over 50 years ago with theestablishment of the Dorena TreeImprovement Center (now DorenaGenetic Resource Center [DGRC],Cottage Grove, Ore.) as headquartersfor the white pine blister rust diseaseresistance program for western whitepine and sugar pine. With an expand-ing focus on improved growth andproductivity in coastal Douglas-fir andother commercial conifer species, theregion also became an early memberof the PNW Progressive Tree Improve-ment Program, which was establishedin 1966. The region’s genetics programwas further expanded throughout the1980s with large investments in seedorchards and progeny test sites for pri-ority species in eastern Oregon andWashington, including ponderosapine, Douglas-fir, western larch, andlodgepole pine.

Fast forward to 2017, a changed land-scape where reforestation needs in theregion are no longer driven primarily bytimber production objectives or regener-ation harvest activities, but rather byresponses to disturbances such as wild-fires, insect and disease outbreaks, andwind storms. Regional genetics pro-grams have responded to these chang-ing conditions by evolving from a focuson tree improvement for enhancedgrowth and yield to today’s broaderemphasis on genetic resource manage-ment, genetic conservation, and ecosys-

tem resiliency andhealth. This work isspread across numer-ous species, expandingin recent years toinclude non-commer-cial native conifer andhardwood tree speciesas well as native shrubs,grasses, and forbs thatare important forrestoration and landmanagement prioritiessuch as creation of fireresistant understoriesand enhancement offorage and habitat forwildlife, including polli-nator species.

Staff and facilities

Geneticists: In addi-tion to a regionalgeneticist, there arefour area geneticists inthe region. Each areageneticist serves multi-ple forests, but alsoprovides specializedexpertise, consultation, and trainingacross the entire region on specific top-ics such as seed biology, cone collec-tion, analysis of genecology studies andprogeny test data, interpretation ofmolecular data, design and implemen-tation of genetic conservation strate-gies, and more. Additional resourcesinclude the DGRC geneticist, as well asthe five-needle pine and Port-Orford-cedar program managers. The regionalgeneticist provides oversight and sup-port to the area geneticists, DGRC, andthe regional Bend Seed Extractory(Bend, Ore.), and also serves as theregional native plant program manager.

Dorena Genetic Resource Center: Thisunique, multi-faceted facility houses aworld-leading applied disease resist-ance testing program for non-nativepathogens including white pine blisterrust and Port-Orford-cedar root disease.Containerized Port-Orford-cedar seedorchards are maintained in DGRCgreenhouses for operational productionof disease resistance seed for reforesta-

tion in southwest Oregon and northernCalifornia coastal forests. Newly upgrad-ed freezer storage facilities house geneticconservation seed collections for at-risktree species, as well as a unique collec-tion of seed from the over 40,000 parenttrees represented in Region 6 progenytest sites and seed orchards. DGRC alsoproduces small quantities of container-ized seedlings for national forests andpartner clients. Over 120 new speciespropagation protocols have been devel-oped through this work. DGRC assists inregional trainings and is home to theUSFS national Tree Climbing Program, acertification program to ensure safe andeffective tree climbing for Forest Serviceand other agency work, including conecollection, red cockaded woodpeckernest placements, snag creation, smokejumper training, and monitoring forAsian long-horned beetle. In 2016,DGRC celebrated its 50 years of accom-plishments and shared with partnerstheir current work and bright vision forthe future.

14 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

Genetic Resource Management in theUSFS, PNW Region: Past, Present, and Future

T

SOURCE: VICKY ERICKSON

USFS seed orchards are positioned in 50 locationsand cover approximately 1,900 acres.

Regional priorities andchallenges

Described below are some high-lights of Regional Genetic ResourceManagement program priorities andaccomplishments, as well as severalinitiatives underway in response tonew challenges such as expandingexotic pests and forest health issues,climate degradation, and the conser-vation of at-risk tree species.

1. Protect and maintain seedorchards,breeding orchards,and genetictest sites: These installations are invalu-able and irreplaceable repositories forgenetic conservation and use in futurebreeding efforts, and for monitoring theimpacts of new insects, pathogens, or achanging climate. They also provide themost efficient and economical source ofhigh-quality and genetically diverse seedfor reforestation. At present, there are 50seed orchard installations totaling over1,900 acres on Oregon and Washingtonnational forests. Significant investmentsare made annually across the region inpriority maintenance activities such asorchard fence repairs and replacement,mowing, and fire protection.

2. Replenish reforestation seed-banks and re-initiate seed trainingprograms: The region’s reforestationseedbanks are aging and rapidlydiminishing, particularly for thosespecies and national forests frequentlyaffected by wildfires. In addition, theregion has lost many seasoned refor-estation specialists experienced inseed use planning, seed biology, andcone collection procedures due toretirements. Consequently, the regionhas re-vamped online and in-personcone and seed training programs andestablished a new regional contract forcone collection. In 2016, Region 6national forests made the largest conecollection in over two decades, withnearly 1,500 bushels of cones (425pounds of cleaned seed) collectedfrom six different species. WashingtonState Department of NaturalResources (WADNR) and several pri-vate companies also took advantage ofthe 2016 bumper crop through per-mits to collect cones in national forestwild stands and seed orchards.

3. Combat invasive pathogens andinsects through development of geneti-cally resistant planting stock for anexpanded array of priority species and

populations: DGRC’s disease resist-ance testing program has grown inrecent years to now include 8 five-needle pine species and geographicsources from the US, Canada, andMexico. One of the more prominentnew focal species is whitebark pine,which has been proposed for federallisting under the Endangered SpeciesAct. Port-Orford-cedar, another focalspecies, involves a vastly smaller geo-graphic zone but is important to USFS,Oregon Department of Forestry, tribes,private landowners (industrial andnon-industrial), NGOs, and the gener-al public. The region has strong work-ing partnerships with these groupsand interest in resistant seed is grow-ing. DGRC’s geneticist also providestechnical assistance in developing pro-grams elsewhere, such as the koa wiltresistance program in Hawaii.

4. Accelerate and expand genetic con-servation and restoration efforts forhighly vulnerable species and popula-tions: Many ecologically important PNWconifer species are in decline due toinsect and disease outbreaks, wildfires,climate change, and other disturbancesand stressors. A recent regional assess-ment indicated that high-elevationspecies and populations in isolatedareas or at the edge of a species distribu-tion were at the highest risk of loss. Theassessment identified priority speciesand geographic areas of special concernand set goals for seed collection andother conservation and restorationwork. International collaborations withthe U.K. Forestry Commission and theMillennium Seed Bank have been lever-aged to help collect seed for genetic con-servation, with seed from over 1,200individuals from seven different speciescollected to date. The collection is storedlocally at DGRC and at the USDANational Center for Genetic ResourcesPreservation (Ft. Collins, CO).

5. Develop climate-based solutionsfor seed deployment: Under any climatescenario, the selection of appropriateseed sources provides the foundationfor successful reforestation. With pre-dicted changes in future climates, themovement of seed sources duringreforestation will become an increas-ingly important strategy for maintain-ing forest productivity and re-aligningspecies and genetic resources to copewith new stressors and site conditions.

In preparing for the future, geneticistsare working with silviculturists andreforestation specialists to evaluateexisting seed inventories and test newmodels such as the Seedlot SelectionTool (see Harrington and St. Clair arti-cle) to identify areas where seed may bewell suited to future climates, or tochoose the best seed source for a par-ticular planting site under predicted cli-mate scenarios. These and other relatedclimate change adaptation activities areguided by a national USFS strategy doc-ument: “Genetic Resource Manage-ment and Climate Change: GeneticOptions for Adapting National Foreststo Climate Change.”

Looking ahead

Maintaining productive andresilient national forests into the futurewill require new tools, practices, andre-focused investments in all areas ofland stewardship, including geneticresource management. Retainingexpertise and a supporting plant pro-duction infrastructure (i.e., nurseries,seed extractories, disease resistancescreening centers, and seed and breed-ing orchards), coupled with strong sup-port from research, management, andour external partners are all vital ele-ments for sustaining a strong and for-ward-thinking PNW Genetic ResourceProgram. Although there are manyconstraints and undoubtedly bumpsahead, the region is well-positioned torespond to the needs and challenges ofthe next 50 years and beyond. ◆

Vicky Erickson is regional geneticistand Native Plant Program manager,US Forest Service, Pacific NorthwestRegion, in Pendleton, Ore. She can bereached at 541-278-3715 or [email protected].

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 15

Tom [email protected]

206 300 9711www.arborinfo.com

Providing information about trees and forests

BY BRAD ST. CLAIR ANDGLENN HOWE

orest genetics research in thePacific Northwest goes back to the

beginning of the 20th century. One ofthe first long-term studies undertakenby the US Forest Service, and one ofthe first forest genetics studies inNorth America, was a provenancestudy of geographical variation andtrait inheritance of Douglas-fir. Thisstudy, called the Douglas-Fir HeredityStudy, was established by Thornton T.Munger in 1915 at the Wind RiverExperiment Station and other sites inOregon and Washington. It was recent-ly remeasured, and is still providinginsights more than 100 years later.Another important study was the WindRiver Arboretum, a study looking atnon-native species for use in thePacific Northwest. Both studies wouldprove to be highly influential on sub-sequent thinking about appropriatespecies and seed sources for reforesta-tion. Meanwhile, poor natural regener-ation after large fires in the early 20thcentury led to an interest in tree plant-ing and the establishment of the WindRiver Nursery in 1910. Early foresters,however, did not pay particular atten-tion to their choice of seed source,resulting in poor growth and survival

of many stands. Experiences withplantation failures and the resultsfrom the species and provenance trialsensured the preference for nativespecies, and led to the development ofseed collection guidelines and the firstdelineation of seed zones in the 1940s.By the 1960s, a seed source certifica-tion system was adopted, and in 1966,seed zone maps for Oregon andWashington were published.

Interest in forest genetics and treeimprovement took off in the 1950s. Anincrease in logging as a result of thepost-war housing boom led to anincreased interest in productionforestry and a shift from largely naturalregeneration to planting. Forestersbecame keenly interested in plantingwell-adapted seed sources and in possi-ble improvements through appliedgenetics. The Douglas-Fir HeredityStudy was often used to demonstratethe importance of genetics by pointingto the large differences in growthamong family row plots. In 1954, theIndustrial Forestry Association (IFA)hired Jack Duffield to begin a programof Douglas-fir tree improvement. Alsoin 1954, Roy Silen established aresearch program on forest geneticsand tree improvement at the USFSPacific Northwest Forest and RangeExperiment Station (now the Pacific

Northwest Research Station {PNWRS}).In the late 1950s, Alan Orr-Ewing beganresearch on tree improvement andgenetic variability of Douglas-fir for theBritish Columbia Forest Service (nowthe BC Ministry of Forests, Lands, andNatural Resource Operations{MFLNRO}). In Idaho, Richard Binghambegan to breed for blister rust resist-ance in western white pine at the USFSIntermountain Forest and RangeExperiment Station (now the RockyMountain Research Station {RMRS}).Meanwhile, the need for an organiza-tion to bring together researchers andpractitioners to share information wasrecognized, and the Western ForestGenetics Association was established in1955. It continues to be the primaryforum for forest genetics in westernNorth America.

Tree improvement programs grewover the next decades, particularly forDouglas-fir. A major obstacle forDouglas-fir, however, was the issue ofgraft incompatibility in clonal seedorchards. In 1966, Roy Silen proposedan alternative strategy to get aroundthis issue by using seedling seedorchards created from full-sib crossesof parents in the field. The parentswould be intensively selected fromprogeny tests of a large number ofopen-pollinated families from nativestands. Under the leadership of RoySilen and Joe Wheat of the IFA, this“progressive” tree improvement pro-gram brought together many publicand private landowners to cooperatein selection and testing. This programlater became the Northwest TreeImprovement Cooperative (NWTIC).

At about the same time, the BritishColumbia Forest Service expanded itswork with a diallel crossing program oftheir Douglas-fir selections followedby field testing. The BC program latercame under the auspices of the ForestGenetics Council. The Inland EmpireTree Improvement Cooperative(IETIC) was founded in 1968 to devel-op improved ponderosa pine in east-ern Washington, northern Idaho, andwestern Montana, and expanded theirprogram in 1974 to include severaladditional native conifer species.Weyerhaeuser initiated an independ-ent research and tree breeding pro-gram. Other independent programswere begun that later came together

16 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

Building on a Century of ForestGenetics Research

F

PHOTO COURTESY OF BRAD ST. CLAIR

An early picture of the Wind River Nursery illustrates the story of reforestingthe burns shown in the background.

under the auspices of the NWTIC.Today, tree improvement programsexist for a wide variety of conifers,including Douglas-fir, western hem-lock, ponderosa pine, interior spruce,western redcedar, lodgepole pine,western larch, Port-Orford-cedar, andfive-needle pines. The University ofWashington and Washington StateUniversity had large research pro-grams aimed at developing hybridpoplars, and several industriallandowners planted extensive poplarplantations in Oregon andWashington. Research on diseaseresistance continues at the USFSDorena Genetic Resources Center,RMRS, and BC MFLNRO.

For most species, geneticallyimproved planting stock is producedin seed orchards. Thus, seed orchardresearch has been an important com-ponent of forest genetics research inthe Pacific Northwest. The problem ofgraft incompatibility in Douglas-firwas mostly solved by Don Copes at thePNWRS through a research and breed-ing program that led to the develop-ment of graft-compatible rootstock.Because of his efforts, clonal seedorchards are the foundation ofDouglas-fir tree improvement, andnearly all seed orchard clones aregrafted onto “Copes’” rootstock.

Other important seed orchardresearch has been conducted by thePNWRS, Pacific Northwest TreeImprovement Research Cooperative(PNWTIRC), BC MFLNRO, andWeyerhaeuser Company. Improve-ments have been made in orcharddesign (spacing and arrangement ofclones), flower stimulation, crownmanagement, supplemental mass pol-lination, pest management, andbloom delay to control pollen contam-ination. Tom Adams at Oregon StateUniversity (OSU) pioneered the use ofallozyme genetic markers to under-stand and manage mating systemsand pollen contamination in seedorchards, and this work was extendedto SSR genetic markers by thePNWTIRC. The PNWTIRC has alsostudied the genetics of important traitssuch as growth, stem form, wood qual-ity, cold hardiness, and drought hardi-ness; developed breeding approachessuch as early selection; and developedgenetic markers that are now being

used by breeders. Approaches foradvanced generation breeding, fieldtesting, and data analysis were devel-oped by Bob Campbell and RandyJohnson at the PNWRS, and later bythe NWTIC.

Other forest genetics researchfocused on population genetics andgeographic variation in adaptive traits.The advent of allozyme genetic mark-ers gave us a better understanding ofgene flow, population structure, andphylogenetics of several species.Genecology research was conductedusing short-term common gardenstudies to understand how seedlingperformance is related to the climaticenvironment of their parents. The pio-neering work of Jerry Rehfeldt at theRMRS, and Bob Campbell and FrankSorensen at the PNWRS, provided anunderstanding of the geographic dis-tribution of adaptive traits that wasimportant for refining seed zones andbreeding zones. This genecologyresearch will become increasinglyimportant as we grapple with climatechange and to help us understandhow forests will respond and what wecan do about it. Recently, the PNWRSestablished a provenance test using areciprocal transplant design in whichseed sources from a wide range of cli-mates were planted at test sites of awide range of climates. Results fromthis study, called the Seed SourceMovement Trial, are being used tosimulate the effects of climate change,and to evaluate options for “assistedmigration”—a strategy that involvesmoving populations to areas wherethey are expected to be better adaptedto future climates.

Finally, we’ve entered a new age—the age of genomics. As in the past,new knowledge, approaches, and toolswill continue to make tree breedingand adaptation to climate changemore effective, less costly, and faster.

We now have complete genomesequences for several important foresttrees, with more to come. Millions ofgenetic markers are now available totree breeders—in contrast to the 20 orso allozyme markers available indecades past. We can now directlystudy the expression of the genes thatare important to adaptation to cli-mate, disease resistance, or increasedgrowth. In the PNW, this work hasbeen active at the University of BritishColumbia, OSU, PNWRS, and RMRS.New approaches for gene editing(using CRISPR) have grabbed theattention of the biological sciencesand the public. Although geneticallymodified trees are unlikely to be wide-ly used in the Pacific Northwest, thesetechnologies and their potential bene-fits are being studied by the TreeBiosafety and Genomics ResearchCooperative at OSU.

The past 100 years has seen enor-mous progress in our understandingof forest genetics and our ability tobreed trees—but we cannot rest. Thefuture will bring even greater chal-lenges, including climate change, theintroduction of new pests, and greaterdemand for natural resources. Thus,we must continue to develop new for-est genetics knowledge, approaches,and tools to help sustain our forestsand society. ◆

Brad St. Clair is research geneticist withthe US Forest Service Pacific NorthwestResearch Station in Corvallis, Ore. AnSAF member, he can be reached at 541-750-7294 or [email protected]. GlennHowe is director of the Pacific NorthwestTree Improvement Research Cooperativeand associate professor in theDepartment of Forest Ecosystems andSociety at Oregon State University inCorvallis. He can be reached at 541-737-9001 or [email protected].

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 17

www.greencrow.com

Certifiably Proudof the Washington

Tree Farm Program

BY GLENN HOWE

he Pacific Northwest Tree Improve-ment Research Cooperative

(PNWTIRC; www.fsl.orst.edu/pnwtirc/)conducts forest genetics and treebreeding research that supports forestmanagement by federal and stateagencies and a multi-billion-dollar for-est products industry. Over the years,our members have included privatecompanies and governmental agen-cies in Oregon, Washington, andBritish Columbia. The focus of ourresearch is on Pacific Northwest treespecies and the development of newtree breeding tools and approaches.The knowledge and tools we developare made available to breeders inreports, scientific publications, andthrough technology transfer meetingsand workshops. Most of our researchattention has been on Douglas-fir, butwe also conduct research on other treespecies such as western hemlock andwestern white pine. The PNWTIRCwas formed in 1983 to address con-cerns that forest genetics research wasnot keeping pace with the rapidexpansion of applied tree breedingprograms. Today, the cooperative has14 member organizations.

Tree breeding research

The initial focus of the PNWTIRCwas to understand the genetics ofDouglas-fir growth and stem formtraits. How can we efficiently andaccurately measure these traits? Arethey heritable? How much genetic gainis possible if these traits are includedin breeding programs? What is theirrelative importance—genetically andeconomically? Results from these earlystudies and subsequent research high-

lighted the importance of stemdefects, particularly the loss in valuecaused by ramicorn branches, whichare large, steep-angled branches thatare often associated with second flush-ing in the summer. These early, short-term studies provided quick results tobreeders because they were conductedin existing genetic tests owned byPNWTIRC members.

The next phase of the PNWTIRCfocused on newly designed experi-ments, such as the early testing study.In this experiment, we grew Douglas-fir seedlings in outdoor nursery bedsand greenhouses, and then comparedtheir performance to siblings that hadalready been growing in the field for 15years. This retrospective approachdemonstrated that we can predictlonger-term field performance bymeasuring families in their first or sec-ond growing season—at least wellenough to practice early culling. Earlyculling is the removal of the poorestperforming families at the seedlingstage, before they are planted in long-term and expensive field tests. Thus,we can use this approach to lower treebreeding costs and increase the qualityof the field tests by allowing us to plantsmaller test plantations that have lessenvironmental variation.

In the 1990s, the PNWTIRC concen-trated on the genetics and physiologyof adaptive traits such as cold hardi-ness and drought hardiness. In keep-ing with the early testing theme, amain goal of this work was to developearly screening protocols for identify-ing cold hardy and drought hardygenotypes. We developed a cold hardi-ness testing protocol that involves col-lecting cuttings from progeny tests andthen testing them using artificial freezetests in programmable freezers. These

artificial freeze tests are now widelyused to improve fall cold hardiness inDouglas-fir breeding programs.

We also developed a seedling testingprotocol for drought hardiness. Thisapproach involves imposing droughtstress in raised nursery beds and thenassessing drought hardiness by meas-uring foliage damage, diameter growth,and the loss of water flow through thestem caused by embolisms. Because ofclimate change, our work on droughthardiness continues with recentlyestablished genetic tests on hot and drysites in southern Oregon.

Research by the PNWTIRC and oth-ers led to the adoption of new acoustictools for measuring and improvingwood stiffness in Douglas-fir breedingprograms. Between 2005 and 2013, weconducted two major studies ofDouglas-fir wood properties. Douglas-fir is renowned for its strong and stiffwood, but breeders are concerned thatwood quality could decline becausefaster-growing trees are being harvest-ed at younger ages. Thus, the aim ofthis research was to develop effectiveand cost-efficient ways to identifygenotypes with stiff wood. Tools thatmeasure acoustic velocity had alreadybeen used to sort mill logs based onwood stiffness. To predict stiffness, onecan strike the log with a hammer andthen measure the speed of the result-ing sound waves. We demonstratedthat these same tools (e.g., Hitman orTreeSonic) can be used on standingtrees or logs thinned from genetic teststo identify superior genotypes inbreeding programs. A core element ofthis project was a study in which weharvested trees from 25-year-oldgenetic tests and then milled nearly400 of them into 2x4s to test our woodstiffness predictions. Later, we extend-ed this research to much younger treesand western hemlock.

Seed orchard research &molecular genetic markers

In the Pacific Northwest, seedorchards are fundamentally importantfor capturing genetic gain and deliver-ing it to the field. Improved genotypesfrom breeding programs are plantedtogether in seed orchards where theycan cross-pollinate to produceimproved seed. Thus, seed orchardresearch has also been an important

18 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

Cooperative Brings Life to TreeBreeding Tools and Approaches

T

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 19

part of our mission. Forexample, we studied seedorchard design and man-agement, including treespacing, crown manage-ment, early flowering,and the use of moleculargenetic markers to under-stand mating patterns inseed orchards. We con-ducted a 15-year study ofminiaturized seedorchards, which are alter-natives to traditional seedorchards where trees areplanted at a close spacingand maintained at aheight of only 10 feet orless. Because of high per-hectare seed yields,miniaturized orchards arepotentially desirable forcapturing genetic gains atan early age. The smallsize of the trees also facilitates con-trolled crossing, supplemental masspollination, overhead irrigation, andcone harvesting. The methods of earlyflower stimulation we developed as partof our miniaturized seed orchard pro-gram are widely used to increase theyields of genetically improved seed inDouglas-fir seed orchards.

In Douglas-fir, we developed genet-ic markers called SSRs that are beingused by tree breeders to identify mis-labeled trees in seed orchards, deter-mine the parents of open-pollinatedseedlots, and measure pollination con-tamination. Pollen contamination,which occurs when trees in nearbystands mate with the genetically supe-rior orchard trees, reduces geneticgains and may lower adaptability. TheDouglas-fir SSRs we developed arebeing used by seed orchard managers,breeders, and other forest geneticists,mostly through the genotyping servic-es offered by the USFS National ForestGenetics Laboratory.

Current focus & future research

Currently, the PNWTIRC is helpingto bring the latest genomic approach-es to tree breeding. New, large-scalegenomic technologies and advancedstatistical approaches have alreadytransformed crop and livestock breed-ing, and are likely to do the same fortree breeding. For example, we used

genomic approaches to identify hun-dreds of thousands of genetic markerscalled SNPs (single nucleotide poly-morphisms) in Douglas-fir. SNP mark-ers are now being used to test anapproach called genomic selection toimprove growth and wood quality inDouglas-fir, and to enhance blisterrust resistance in western white pine.These topics are discussed in greaterdetail in the article on forest genomicsand biotechnology.

The fundamental mission of thePNWTIRC has been the same since1983—to advance tree breeding andour understanding of native tree pop-ulations in the Pacific Northwest. Theprospects for using genomics toenhance tree breeding are great andwill be a key area of research into thefuture. At the same time, climatechange imposes challenges to treebreeders and forest managers. Thus, itis particularly important to under-stand how trees are genetically adapt-ed to climate. Future research by thePNWTIRC will continue to addresstopics such as seed zones and breed-ing zones, assisted migration, and thegenetics of adaptation to cold anddrought. For example, we are collabo-rating with the US Forest Service andthe Conservation Biology Institute todevelop a suite of tools that can beused to implement forest managementpractices designed to help forests

adapt to climate change.The first of these tools is aweb application calledthe Seedlot Selection Tool(SST; https://seedlotselectiontool.org/sst/).The SST is a GIS mappingtool designed to help for-est managers match seed-lots with planting sitesbased on selected climatechange scenarios.

As a research coopera-tive, collaboration, team-work, and the sharing ofknowledge and resourcesare fundamental to whatwe do. Members of thePNWTIRC not only pro-vide funds needed to con-duct PNWTIRC research,but also field sites, plantmaterials, and the expert-ise and energy needed to

make the research possible and rele-vant. Additionally, we have many out-side partners and funding organizationsthat have contributed to our success. ◆

Glenn Howe is director of the PacificNorthwest Tree Improvement ResearchCooperative and associate professor inthe Department of Forest Ecosystemsand Society at Oregon State Universityin Corvallis. He can be reached at541-737-9001 or [email protected].

PHOTO COURTESY OF GLENN HOWE

The PNWTIRC conducts research on tree breeding methods andthe production of genetically improved seed. This photo showsJim Smith next to grafted trees that were part of a 15-year studyon the establishment and management of Douglas-fir miniatur-ized seed orchards.

Restore ourfederal forests

to restore ourrural communities

Join us @HealthyForests.Org

BY SARA LIPOW

vital compo-nent of forestry

today for most com-mercially importanttree species is refor-estation with geneti-cally improved plant-ing stock producedby tree improvement. Tree improve-ment is the process by which knowl-edge of tree genetics is coupled withknowledge of silviculture, economics,and wood science to select and refor-est with trees that possess desirabletraits. These desirable traits usuallyrelate to increased wood yield or high-er wood quality, which are the focus ofthis article. Other desirable traits suchas disease resistance and forest healthare discussed elsewhere in this issue.

Tree improvement is expensiveand takes decades

Foresters are accustomed to long-term thinking. During reforestation, wemake investments in site preparationand tree planting knowing that finan-cial gain from these activities will notbe realized until trees are harvesteddecades later. Tree improvement alsotakes decades, and its investment startslong before a tree is even planted.

Most tree improvement programsin western North America began in the1950s to 1980s and followed a similarpattern. Trees growing in naturalstands were identified and cones werecollected from them. Sometimes these“parent trees” were selected after com-paring them to other trees in a uni-form stand, but sometimes they weresimply good looking trees that were

easy to collect cones from. Regardless,seeds from those parent tree cones(their progeny) were then grown infield trials, called progeny trials. Thesefield trials were expensive: they werereplicated across multiple sites witheach tree individually tagged andmeasured several times. By measuringa large number of progeny from eachparent tree, geneticists identified theparent trees that possessed desirabletraits due to having desirable genes.Traits of interest, especially height anddiameter, were typically measured atleast until 15 years of age and some-times longer.

This first-generation tree improve-ment testing allowed geneticists toidentify families of trees that displayeddesirable traits that were inherited andthus could be passed on. For mostspecies, the best trees, either progenyor parent trees, were then grafted intoa seed orchard and managed to pro-duce seeds for reforestation. Seedorchards are managed more like fruitorchards than like forestry plantations;they are mowed, sometimes irrigated,and often treated with plant hormonesor other methods to stimulate coneproduction. They are expensive toestablish and maintain. The time ittakes to produce seed from a seedorchard tree varies, but generallyranges from eight to 15 years forspecies such as Douglas-fir, ponderosapine, and noble fir. Thus, productionof genetically improved seed involvesyears of expenses for both treeimprovement research and for seedorchard management. In addition, forsome species, tree improvement hasgone on to a second generation wherethe best trees from the first generationwere crossed together, seeds from

those crosses grown in another roundof replicated progeny trials, and thebest second-generation trees select-ed—again a decade’s long process. Fora few species, including coastalDouglas-fir and western hemlock, athird generation of tree improvementis in progress.

Genetic gain

Most genetically improvedseedlings planted in western NorthAmerica during 2017 will be fromsome type of first-generation seedorchard. Those seedlings were pro-duced by trees or their progeny thatwere found growing naturally in theforest—but identified through treeimprovement as having desirabletraits—and then raised in a seedorchard where they were pollinated byother, similarly selected trees. Givenhow “natural” these seedlings are, howmuch better are they compared toseedlings produced by “woodsrun”seed collected from natural stands? Inother words, how much genetic gaindo they achieve? The answer dependson many factors, especially the num-ber of parent trees tested and the qual-ity of the progeny testing.

The company I work for, RoseburgForest Products Company, participatesin tree improvement cooperatives forDouglas-fir that give us access to thou-sands of first-generation parent treesappropriate to our Oregon land base,and we have placed the top ones inintensively managed seed orchards. Asa result, the first-generation seedlingswe plant are expected, on average, toproduce plantations that at age-15 are8-15% taller with 15-40% more volumethan if we had used woodsrun seed.These values are expressed for planta-tions that are 15 years old, which was acommon age for the final measure-ment of many of the first-generationtests. All this genetic gain will not berecovered at rotation. Based on predic-tions from growth models and compar-isons from other species, I conserva-tively estimate that half of this geneticgain will persist to rotation. For someof our land base, we have begun plant-ing second-generation seed, which willyield even higher genetic gains.

Roseburg is not alone in achievingthese high genetic gains. Similar valuesare expected for most landowners in

20 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

Genetic Gain and the EconomicBenefits of Tree Improvement

A

Connecting Forest Landowners withSeedlings, Services and Contractors

DISCOVER Our Interactive Websitewww.forestseedlingnetwork.com

BUY/SELL SEEDLINGS • FIND VENDOR SERVICES & CONTRACTORS • VALUABLE RESOURCES

western Oregon and Washington thathave invested in cooperative treeimprovement for Douglas-fir andwestern hemlock. Genetic gains areusually lower for minor species, suchas noble fir, which have lower invest-ments in tree improvement.

A high return on investment

When many plantations are estab-lished using genetically improvedstock, the increased wood productioncan be substantial, and this provides astrong economic motivation forinvestment in tree improvement pro-grams. The return on investment canbe quantified using discounted cashflow analysis: Landowners input treeimprovement and seed orchard costs,when the costs were incurred, acreageto be planted, and the genetic gainsachieved. Then, using a forest growthmodel, the amount of wood productsproduced over time using geneticallyimproved seed versus woodsrun seedcan be estimated and a monetaryvalue assigned to those wood prod-ucts. A “discount rate” (sometimescalled the compounding interest rate)is included in the analysis to accountfor the time value of money, since

money tied up for decades in treeimprovement is not available for otherinvestments. With this approach, pro-jections of the future value of woodproducts can be discounted to arriveat a present value estimate for invest-ing in tree improvement and seedorchards. Such economic analysesdone for many tree improvement andseed orchards programs in westernNorth America demonstrate a highreturn on investment under reasonablegenetic and economic assumptions.

The economic benefits from treeimprovement are realized in severalways. Plantations established withgenetically improved stock have higheryields. They often also produce highervalue wood products. Additionally,since they grow substantially faster,the wood can be harvested sooner,reducing the rotation age, and allow-ing the forestland to be cycled morequickly; this is captured in economicterms as increased land expectationvalue. Genetically improved stock canalso result in more uniform stands,leading to lower harvesting and trans-portation costs. Inflation is also a con-sideration: when wood products expe-

rience positive real rates of inflationcompared to other goods in the econ-omy, this magnifies the genetic gainsin yield and quality, making invest-ment in tree improvement even morefavorable.

The return on investment from atree improvement and seed orchardprogram is tied to the size of the landbase it serves. Whereas the costs ofmost silvicultural treatments areincurred on a per acre basis, (e.g., veg-etation control and fertilization), manycosts associated with tree improve-ment including selection, breeding,and testing occur at the program level.Therefore, landowners receive higherreturns as they reforest more acreswith genetically improved stock. Thiseconomy of scale has resulted in theformation of tree improvement coop-eratives and accounts for their remark-able persistence despite many changesin land ownership, and thus coopera-tive membership. Moreover, geneticgains are cumulative across genera-tions of tree improvement and acrossrotations of a plantation. Thus, where-as other silvicultural treatment mustbe repeated with each rotation, it isnot necessary to repeat the costs of anentire tree improvement cycle for eachrotation. For these reasons, continuedinvestment in tree improvement andseed orchards, and continued eco-nomic benefits to its participants andthe regional economy, is anticipatedfor valuable plantation species inwestern North America. ◆

Sara Lipow is a Tree Improvementsupervisor with Roseburg ForestProducts Company in Lebanon, Ore.She can be reached at 541-259-2651 [email protected].

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 21

PHOTO COURTESY OF KEITH JAYAWICKRAMA

Realized genetic gain trials that compare plots established with woodsrunstock (right) to those established with genetically improved stock (left) helpdemonstrate to forest managers the economic benefits of tree improvement.The faster growth rate achieved by tree improvement is visually apparent andstatistically significant in the Molalla realized genetic gain trial for Douglas-fir,established in 1997 by the Northwest Tree Improvement Cooperative.

FORESTR 4ESTMGRFORESTR

4ESTMGR

HOPKINS FORESTRYForest Managers performing herbicideapplication, young stand management,

harvest management, contract compliance,inventories, and forestry/natural

resources education

Dick & Paula Hopkins360-492-5441

[email protected]

BY CONSTANCE A. HARRINGTON AND BRAD ST. CLAIR

limate change inthe 21st century

is likely to dramatical-ly alter the growingconditions that PacificNorthwest treespecies experience. Ithas been suggestedthat foresters plan forthese changes bymoving seed sourcesto locations where theseed-source environ-ment and the futureclimate will be simi-lar. Some people havecalled this type ofseed-source move-ment “assistedmigration” with the idea that we arehelping the plants move to better suitedsites faster than they would naturally.But it is important to realize that peoplehave moved seed sources to new loca-tions for centuries without using thisterm. Think of David Douglas, the earlybotanist, plant collector, and the onefor whom Douglas-fir is named, whosent thousands of seeds back to theBritish Isles for propagation.

But when it comes to establishingforest stands, rather than ornamentalplants or small plantings of exoticspecies, foresters wonder how far itwould be safe to move trees. Should weselect seed lots most similar to the cur-rent climate, or the climate in 20 years,or the climate in 40 years? After all, ifthe climate hasn’t changed much yet,what would be the benefits or risks inplanting seed lots which were adaptedto a different climate (say one warmeror drier than the one at the outplantingsite) as opposed to local sources?

In 2009, we put out a call to landmanagers to work with us on a newresearch trial designed to help answerquestions about seed-source move-ment by providing combinations ofseed sources and test sites where wecan evaluate various aspects of plant

adaptation. We received a greatresponse and currently have 10 organi-zations involved: Bureau of LandManagement, Cascade TimberConsulting, Giustina Land and Timber,Hancock Forest Resources, Lone RockTimber Company, Port Blakely TreeFarms, Roseburg Resources, StarkerForests, USDA Forest Service StoneNursery, and Washington Departmentof Natural Resources. In addition, wehave students and faculty from OregonState University, Evergreen StateCollege, and the University of BritishColumbia working with us.

We collected from 60 diverse popu-lations selected from natural stands inCalifornia, Oregon, and Washington(see Figure 1); these were from coastaland inland sites and represented arange in elevations. We planted theseedlings on nine sites in Washingtonand Oregon in the fall of 2008 or springof 2009. All the sites were fenced to pre-vent browsing by deer and elk and weretreated operationally (site preparationor vegetation control treatments) byour partners prior to or after planting.

The seed sources cover 12 geograph-ic regions: three in California, three insouthern Oregon, three in central

22 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

The Douglas-fir Seed-Source MovementTrial Yields Early Results

C

ConstanceHarrington

Brad St. Clair

SOURCE: C. HARRINGTON

Figure 1. Locations of the seedsources and outplanting sites in theDouglas-fir Seed-Source MovementTrial.

SOURCE: C. HARRINGTON

Figure 2. Height by geographic source eight years after planting at theNortons site on Starker Forests land west of Philomath, Ore. In general,trees from the coastal families in each latitudinal band grew better thanthose from inland low elevations sources or high elevation sources.

Oregon, and three in Washington. Theoutplanting sites along three generallatitudinal bands include a coastal site,a low-elevation inland site, and a higherelevation inland site. Each site includes48 plots—four blocks with each of the12 regions planted in each block. Eachsite has a weather station and someadditional environmental sensors.

The sites have been visited manytimes for different types of assessments.We measure the usual things like heightand diameter, and record mortality andtree condition. We have also assessedthe timing of vegetative budburst in thespring and the presence of foliage dis-eases such as Swiss needle cast andRhabdocline on all sites. And on somesites we have collected additional infor-mation related to cold or drought har-diness, flowering, and timing of heightand diameter growth.

So, what have we learned?First, the most important thing we

have learned (or reinforced from othertrials) is that we need to focus on “cli-mate space” and not “geographic space.”That is, it is important to specify how farwe move something in terms of the dif-ferences between the climate at the seedsource site and the climate at the plant-ing site and not in terms of miles thecrow flew or the mountain goat climbedin elevation. The most important vari-ables describe how cold the winters are(e.g., minimum winter temperature) andhow dry the summers are (e.g., summerprecipitation or some measure of evapo-transpiration or aridity). We can movetrees fairly long distances as long as theclimate variables don’t change much (forexamples on how to use this concept toselect seedlots to plant or where todeploy seedlots you have, check out theSeedlot Selection Tool onlinehttps://seedlotselectiontool.org/). Bereassured, we are not suggesting movingtrees from the California coast to theWashington Cascades, but rather youmight consider some small seedlotmovements in a climate-smart manner.Moving trees too far can result in mortal-ity, top dieback, and increased suscepti-bility to some insects and diseases.

Plants from coastal environmentsgenerally are faster growers than thosefrom inland sources, and those fromhigher elevation sources grow lessthan those from lower elevationsources (see Figure 2). This conclusion

has been reported many times fromother studies, but it’s good to see itrepeated in our current trial.

Sometimes mild sites are not thebest. For example, the sites that hadcool weather in the fall had less colddamage to artificial cold events thanthe site which had mild weather in thefall as trees on the milder site did notget the “slow down” signals of tempera-tures close to freezing before they expe-rienced a freeze event.

It has been reported for several horti-cultural plants that warmer winters

result in earlier dates of flowering orbudburst. However, coastal sites insouthern Oregon and California do notcurrently experience many chillinghours in the winter and Douglas-fir hasan obligate chilling requirement; thus,warmer winters in the future on siteswith currently mild winters could resultin no advancement in date of budburstor even delays in budburst as the treeswouldn’t experience enough cool tem-peratures to burst bud promptly in thespring.

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 23

Early Provenance Trials Leadto Contrasting Conclusions

BY KEITH JS JAYAWICKRAMA

Compared to Europe, where many resources were invested in studyingprovenance variation in forest trees, little was done in the Pacific Northwestthrough the heyday of forest tree improvement in the 20th century. Just twoDouglas-fir trials were established, one trial for ponderosa pine was estab-lished, and none for western hemlock, western redcedar or noble fir west of theCascades in Oregon and Washington. The low emphasis on provenance trialswas partly due to lack of interest in importing and testing species from outsidethe PNW, and partly due to a strong commitment on the part of foresters andtree improvers to emphasize local seed sources.

The 1912 Douglas-fir Heredity Study was one of the first forestry provenancetests in the USA, and was established by Thornton T. Munger. In 1915 and 1916,progeny from 120 parents from 13 provenances were planted at six sites.Measurements were taken over the next 100 years (with credit for later meas-urements to Roy Silen); the trial was last measured in 2013. The original objec-tives were to determine the best type of tree to be used for seed collections forartificial reforestation and for seed trees, as well as to determine the influenceof provenance on tree growth. Munger was primarily interested in how progenydiffered between sound versus infected, old versus young, and low- versushigh-elevation parents; thus, the study became known as the Heredity Study,although later emphasis switched more to provenance variation. The trial wasused in the 1960s to make the case for a large-scale tree improvement programfor Douglas-fir and to argue for small breeding zones for this species.

The other Douglas-fir trial was coordinated by Kim Ching at Oregon StateUniversity. Seeds were collected in 1954-1956 from 16 provenances from south-ern Oregon to northern Vancouver Island. Fourteen to 89 parent trees were cho-sen at random at each location for seed collection. The 16 seed lots were out-planted in 1959 on 16 sites (located in the vicinity of seed collections) in a recip-rocal design where one provenance is native to each site. Each planting site waswithin a 40 km radius and 60 m in elevation of a seed collection site. Many siteswere abandoned due to damage caused by frost, browse, drought, and fire. Thesites were measured on a regular schedule through age 25. Close to rotation (46-52 years from seed), total height and DBH were measured at six of the 16 sites(on 7,600 trees). Trial data were used for many publications; in contrast to theDouglas-fir Heredity Study, the case for small seed zones was not compelling. ◆

Keith JS Jayawickrama is director of the Northwest Tree ImprovementCooperative at the College of Forestry, Oregon State University, Corvallis. Hecan be reached at 541-737-8432 or [email protected].

(CONTINUED ON NEXT PAGE)

Our two dry sites in southern Oregon did not have needle diseaseswhen surveyed in 2015. On the other seven sites, all the trees had Swissneedle cast—but some seed sources seemed to tolerate it and grew well. Onthe other hand, there was a big difference in the frequency of Rhabdocline(another needle cast disease) by seed source with the sources which burstbud early most impacted by Rhabdocline. We suspect that the disadvantageof having new foliage available during the time period when many of thefungal spores are dispersing could be a reason why trees from some cool,wet geographic areas—such as the Washington coast—have evolved toburst bud late in the spring.

We learned different thingseach year. For example, thevery hot early summer in2015 changed the pattern ofdiameter growth (see Figure3). In most years diametergrowth occurs throughout theyear from spring through fall.But in 2015 at our J. HerbertStone Nursery site nearMedford, Ore., many of thetrees that were hosting ourelectronic dendrometersshowed the trees stoppeddiameter growth in late Juneand didn’t start again until thefollowing spring. This wouldhave not only reduced thetotal amount of diametergrowth for the year, but alsoreduced the percentage late-wood produced (which couldimpact drought hardiness and wood quality). Sometimes people ask us whatfuture date or climate projection we would suggest they use to plan for thefuture. An easy answer at this point is to suggest you plan for the conditionswe experienced in 2015 (no models needed!).

We also learned that these sites are great for field visits and tours—andnot just ones led by scientists. The differences in survival, growth, or suscep-tibility to insects or diseases are striking and both we and our collaboratorshave used these sites for many formal tours and informal visits. Foresterscan take interested folks (shareholders and board members from the partici-pating companies, regulators, other researchers, and natural resource pro-fessionals) to the sites and SHOW them why we do this and related researchprojects on evaluating and moving seed sources and understanding treeresponses to their growing environments. A picture may be worth a thou-sand words, but a field demonstration is worth a whole lot more than that!

We plan to continue this study for as long as our cooperators will host oursites. The specific types of factors we can study will change over time. Wehave enjoyed taking detailed measurements on young trees, but also lookforward to taking measurements on older trees of other variables thatforesters are also interested in (such as branching, wood quality, stem taper).If you would be interested in working with us, or just in suggesting topics wemight consider for evaluation in the future, please contact either of us. ◆

Constance (Connie) A. Harrington is a research forester and tree physiolo-gist in Olympia, Wash., and Brad St. Clair is a research geneticist inCorvallis, Ore.; both work for the USFS Pacific Northwest Research Station.Connie can be reached at 360-753-7670 or [email protected]. Brad canbe reached at 541-750-7294 or bstclair@ fs.fed.us. Connie and Brad areboth SAF members.

Tracking Changes throughTime-lapse Cameras

Time-lapse cameras can be used todocument differences in budburst andthe timing of height growth. In this exam-ple, we followed trees from two seedsources (southern Oregon Coast andWashington Coast) planted in theBuckhorn2 common garden outplantingsite on Port Blakely Tree Farms land nearChehalis, Wash. In this example, the treeon the left started growing much earlierin the year than the one on the right, butthe one on the right caught up later inthe year. However, which source will dobest in a particular year on a specific sitewould depend on the seed source and theweather conditions during the spring andsummer. A video clip of the full season oftime-lapse images is available athttp://bit.ly/2ev7xbr. ◆

24 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

SOURCE: C. HARRINGTON

Figure 3. Diameter growth at somesites was monitored hourly usingelectronic dendrometers. In 2011, afairly typical year, diameter growth onour outplanting site near Medford, Ore.,started in late March or early April andcontinued until late October or earlyNovember. In 2015, a year with a warmspring and hot summer, diameter growthon that same site started in early Marchand ceased for the season by July 1.

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 25

Access, Easements, Rights-of-Wayand Timber Trespass: What EveryForest Manager Needs to Know,Sept. 20, Olympia, WA. Contact: WFCA.

OSAF Science Workshop, Oct. 3,Linn County Fairgrounds, Albany, OR.Contact: Noelle Arena, [email protected], www.oregon.forestry.org/content/2017-fire-water-health-workshop.

CESCL: Erosion and SedimentControl Lead Training 2-Day inOregon, Oct. 9-10, Eugene, OR. Contact:NWETC.

Seedling Success in the Field:Linking Nursery and OutplantingPractices, Oct. 11-12, Corvallis, OR.Contact: WFCA.

The Hagenstein Lectures—Emerging Voices in Forestry,Oct. 15, World Forestry Center, Portland,OR. Contact: Rick Zenn, [email protected], www.hagensteinlectures.org.

Forest Carnivores and TheirHabitats: A Focus on Fisher,Marten, and Fox, Oct. 19, Linn CountyFair and Expo Center, Albany, OR.Contact: Julie Woodward, 503-807-1614,[email protected], ofricarnivore2017.eventbrite.com.

6th Field Technology Conference,Nov. 7-8, Portland, OR. Contact: WFCA.

2017 SAF National Convention,Nov. 15-19, Albuquerque, New Mexico.Contact: www.eforester.org/safconvention.

Environmental Forensics-SiteCharacterization and Remediation,Dec. 5-6, Tigard, OR. Contact: NWETC.

Mapping the Course: Timberlands,Forest Products Processing andFiber Issues for 2018, Jan. 24-25,Vancouver, WA. Contact: WFCA.

Foresters Forum, Feb. 7-9, Coeurd'Alene, ID. Contact: [email protected], www.forestersforum.com/.

Forest Health 2018, Feb. 28-Mar. 1,OSU Alumni Center, Corvallis, OR.Contact: http://blogs.oregonstate.edu/2018foresthealth/.

OSAF annual meeting, April 18-20,Bend, OR. Contact: Ron Boldenow,[email protected].

WSSAF annual meeting, May 2-4,Longview, WA. Contact: Ellie Lathrop,[email protected].

Calendar of Events

Contact Information

NWETC: Northwest EnvironmentalTraining Center, 1445 NW Mall St., Suite4, Issaquah, WA 98027, 425-270-3274,https://nwetc.org.

WFCA: Western Forestry andConservation Association, 4033 SWCanyon Rd., Portland, OR 97221, 503-226-4562, [email protected],www.westernforestry.org.

Send calendar items to the editor [email protected].

We Remember

Con Schallau1931-2017

Con Schallau was born and raisedin Iowa. He died peacefully in Oregonon July 6 after a long struggle withdementia. He met his beloved wife of60 years, Leanah, while playing in theIowa State Orchestra; him, the oboeand her the violin.

He was kind, thoughtful, andalways greeted others with his won-derful smile. While growing up in theMidwest, his dream was to raise afamily amongst the tall trees andrugged mountains of the west. Out ofcollege, he began cruising timber inthe forests of northern Michigan. Hiscareer as a research forest economistspanned 30 years from Portland toWashington, D.C.

When he retired, he and Leanahvolunteered together at the variousPresbyterian conference centersaround the country, from Sitka, Alaska,to Stoney Point, New York.

They finally settled in Spokane,where they continued their volunteer-ing together at Deaconess Hospitaland the Ronald McDonald House.They were also active in the FirstPresbyterian Church. Con was an SAFmember for 63 years and was electedan SAF Fellow in 1989.

Con was 85 and was preceded indeath by his wife Leanah and siblingsJohn, Wilma, Hollis, and Kay. He is sur-vived by his children Pam and Dan,grandchildren Hannah and Consei,and numerous nieces and nephews.

He was a veteran of the USAF. Anymemorials in his honor may be sent toFirst Presbyterian Church, 318 S.Cedar St., Spokane, WA 99201. ◆

Land Management | Real Estate Services | Wildlife ManagementRecreational Leases | Technical and Data Services

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BY GLENN HOWE ANDSTEVE STRAUSS

orest biotechnol-ogy refers to the

use of biologicalprocesses at themolecular or cellularlevel to make prod-ucts or technologiesuseful to society.Here, we discuss theuse of forest biotech-nology to enhancetree breeding, focus-ing on moleculargenetic markers,gene discovery, andgenetic engineering.

Molecular geneticmarkers have beenused for at least two decades toenhance the production of geneticallyimproved seed. Most molecular geneticmarkers are proteins (e.g., allozymes) orDNA segments that are used to trackthe inheritance of particular chromo-somal locations. Markers can be usedto simply determine how trees arerelated to one another, or to track spe-cific genes. In the 1980s, allozymemarkers began to be used to: (1) iden-tify mislabeled trees in breeding popu-lations and seed orchards; (2) deter-mine the parents of open-pollinatedseedlots; and (3) manage pollen con-tamination in seed orchards. Later, thePacific Northwest Tree ImprovementResearch Cooperative (PNWTIRC)developed simple sequence repeat(SSR) markers for Douglas-fir that arenow widely used by breeders. SSRslargely replaced allozymes becausethey are easier to measure and aremore accurate and precise for measur-ing genetic differences. Recently, thePNWTIRC developed hundreds ofthousands of SNP (single nucleotidepolymorphism) markers for Douglas-fir. Because SNPs are single-letterchanges in the DNA code, they areeasy to measure using automatedtechnologies, and there is a vast num-ber of them in the genome. SNPs have

also been developed for interiorspruce and lodgepole pine by theUniversity of British Columbia’sAdapTree project.

SNP markers are particularly goodfor a breeding approach called genom-ic selection, where superior genotypesare identified by using thousands ofSNP markers per tree. First, we con-struct a prediction equation using atraining population of trees that hasboth SNP and trait data. Then, weapply the prediction equation to otherpopulations (e.g., seedlings) where thetraits have not been measured. Thus,instead of selecting trees based onmeasuring the traits directly, we canuse SNPs. Eventually, it should becomepossible to use genomic selection toreduce field testing, select for maturetree traits (e.g., wood quality) at theseedling stage, and speed geneticimprovement by reducing the lengthof the breeding cycle. While cost is stilla key constraint, the cost of using DNAmarkers continues to decline becauseof advances in technology. In Douglas-

fir, genomic selection is being activelystudied by the PNWTIRC and NorthwestTree Improvement Cooperative usingfunding from the USDA NorthwestAdvanced Renewables Alliance.

Sometimes it is important to knowabout specific genes. For example, sin-gle genes with large effects are occa-sionally found in native tree popula-tions, and it is valuable to know exactlywhich genes these are. Once the geneshave been discovered, genetic markersfor these genes can be developed andused to guide tree breeding. Thisapproach is being taken to developSNP markers for the Cr1 gene in sugarpine, a gene that confers resistance towhite pine blister rust. Knowledgeabout single genes can also be used toalter the characteristics of trees usinggenetic engineering. This can beaccomplished by inserting genes fromother species, or by changing theexpression of native genes in the targetspecies.

Two approaches are commonlyused to discover genes of interest. The

26 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

Biotechnology Research is DevelopingNew Tools for Tree Breeders

F

PHOTO COURTESY OF ANNA MAGNUSON

These eight-year-old poplars, genetically engineered for containment, areundergoing testing in a field trial at OSU. For scale, shown is six-foot-tallThomas Howe.

Glenn Howe

Steve Strauss

first approach is to look for correla-tions between gene expression and thetrait of interest. The second approachis to look for genetic cosegregationusing a genome-wide associationstudy (GWAS). Using GWAS, we cantest many thousands of genes to seewhich are inherited with particulartraits from one generation to the next.

The first step in the pathway fromgene to phenotype (the observablecharacteristics of a tree) is the synthe-sis of messenger RNA (mRNA). If wefind a strong relationship between theamount of mRNA in a tree and a par-ticular trait, the underlying gene islikely to be at least partly responsiblefor the resulting phenotype. Advancedtechnologies for mRNA sequencingnow make it possible to conduct thesestudies on a genome-wide scale, usinga method called transcriptomics. Treesamples are collected from particulartissues at specific times, mRNAs areisolated in the laboratory, the amountsof specific mRNAs are determinedusing sequencing machines, and theunderlying genes are identified. Weused this approach to discover genesassociated with cycles of growth anddormancy in black cottonwood, andscientists at the USFS Pacific North-west Research Station used a similarapproach to study adaptation inDouglas-fir. The genes that we discov-ered are good candidates for influenc-ing traits important to tree breeders,including genes associated with coldhardiness and drought hardiness.

Similar transcriptome studies bythe Tree Biosafety and GenomicsResearch Cooperative (TBGRC) wereused to identify genes involved inflowering, pollen formation, and seedproduction. The TBGRC used RNAsequencing to identify genes expressedin the developing flowers, pollen, andfruits of poplar and eucalypt trees. Thetranscriptome-identified genes arenow being studied using genetic engi-neering.

A major TBGRC project is to devel-op reliable methods for preventingthe spread of exotic and geneticallyengineered trees when required byregulations or for social license. Twohighly effective genetic engineeringmethods are being studied, RNAinterference (RNAi) and gene editingusing CRISPR-Cas9 nucleases. Both

methods are also widely used in med-icine and agriculture, and are highlyspecific and efficient. RNAi poplarshave been produced and found to behealthy and fully sterile in field trials.Gene edited trees have been pro-duced in both poplar and eucalypts,and are currently being studied in thegreenhouse.

A current effort in the forestbiotechnology laboratory at OSU,funded by a major grant from theNational Science Foundation (NSF), isto use GWAS to identify the genes thatcontrol capacity for genetic engineer-ing in black cottonwoods. Many treespecies and genotypes, even withinpoplars, are very difficult to genetical-ly engineer, and the reasons are large-ly unknown. By identifying the genesinvolved, researchers will gain newinsights into the nature of the con-straints and produce new “genereagents” that could be used to over-come barriers in poplar or otherspecies.

TBGRC and forest biotechnologystaff are also active in education

about GMOs in general, and geneticengineering of trees in particular. Aspart of the new NSF grant, we aredeveloping new methods for teachinghigh school students and teachersabout genetic engineering methodsand impacts. This work, broader out-reach efforts, and stewardship tech-nology such as genetic containmentmethods, are essential to develop thesocial license to use genetic engineer-ing in forestry. ◆

Glenn Howe is director of the PacificNorthwest Tree Improvement ResearchCooperative and associate professor inthe Department of Forest Ecosystemsand Society at Oregon State Universityin Corvallis. He can be reached at541-737-9001 or glenn.howe@ oregon-state.edu. Steve Strauss is director of theTree Biosafety and Genomics ResearchCooperative and professor in theDepartment of Forest Ecosystems andSociety at Oregon State University inCorvallis. He can be reached at 541-737-6578 or [email protected].

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 27

SAF National ConventionComing to Portland in 2018

Mark your calendar for October 3-7, 2018, when thelargest gathering of foresters in the country will convergeon Portland for the annual SAF national convention.

While the 2017 national convention in Albuquerque isright around the corner, Northwest members should beaware of this national convention planning effort as manymay be interested in volunteering and attending while it’sin our backyard.

Tammy Cushing is the general chair for convention.Volunteer opportunities will be available before and duringthe convention. From organizing field trips to staffing the raffle, there willbe chances for members to contribute. If you are interested in volunteering,send an email to Tammy at [email protected]. Volunteerservice is a great way to meet people and show Northwest hospitality.

Convention typically draws over 1,600 professionals and students, andwe are expecting a larger crowd in Portland. Foresters and natural resourceprofessionals gather at this meeting to hear the latest research results, net-work, learn about local forest management and practices, get outside andattend field trips, present posters, chat with vendors showing new technolo-gies and services, gain continuing education credits, and collaborate for thefuture.

Be sure to take advantage of the convention being in the Northwest andlet’s show forestry professionals from around the nation our amazing forestresources. Additional information will be shared as we get closer to theevent. ◆

Tammy Cushing

28 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

BY RICHARD SNIEZKO

ome of our native tree species areunder attack, and some face a per-

ilous future. Forest pathogens andinsects are causing high levels of dam-age or mortality that imperil the com-mercial viability of some species or theecosystem function and biodiversity ofour natural forests and urban forests. Inmany cases, the “villains” are non-native pathogens (and the associateddiseases that they cause) or non-nativeinsects; in other cases, a changing cli-mate, change in forest management, orunknown factors have altered the bal-ance in favor of a native pathogen orinsect. Land managers around theworld are facing these issues and areasking for solutions. A potential naturalsolution is to utilize the natural geneticvariation in our native species to helpcounter the damage by these pathogensand insects.

Genetic variation within a species isa key to its potential to evolve in theface of biotic and abiotic threats,including threats caused by pathogensand insects in the cases where theycan’t be eradicated or controlled byother means. In the case of insect ordisease threats, it generally comesdown to genetic resistance of ournative tree species and how we canuse that resistance to keep forestshealthy or restore unhealthy forests.

Basic questions arise such as: (1) Is there genetic resistance?(2) What is the level of resistance?(3) What types of resistance are there?(4) What is the frequency of resist-

ance across the range of the species?(5) Is the level of resistance suffi-

cient for immediate use or will breed-ing be required?

(6) Is the resistance durable (will itlast for decades or hundreds of years)?

(7) What is the timeline and effortneeded to make use of the geneticresistance?

In the cases discussed here, we arelooking only at resistance already pres-ent in our native species and the use

of classical selective breeding, not thepotential for genetically engineeredresistance.

In the Pacific Northwest, white pineblister rust (caused by the fungusCronartium ribicola) and Port-Orford-cedar root disease (caused byPhytophthora lateralis pathogen) aretwo of the most well-known examplesof non-native forest tree diseases thathave caused significant mortality inour forests. All eight species of five-needle pine native to the western USare highly susceptible to white pineblister rust (WPBR). Port-Orford-cedaris the only forest tree species currentlyknown to be highly susceptible to P.lateralis. Damage to spruce by thenative white pine weevil (Pissodes stro-bi) is a notable example of insect dam-age that greatly impairs the use ofSitka spruce and interior spruce inreforestation. Fortunately, there issome genetic variation in our nativetree species and breeding programsare underway in the US and BritishColumbia.

Breeding programs to develop blis-ter rust resistance in sugar pine andwestern white pine started more than

50 years ago by the US Forest Service(USFS), with more recent efforts inwestern white pine in British Columbia.There are USFS regional resistance pro-grams for the Pacific Northwest, PacificSouthwest, and Interior West (with rustscreening facilities at Dorena GeneticResource Center in Cottage Grove,Coeur d’Alene Nursery, and PlacervilleNursery).

In many areas, land managers arereluctant to plant these species with-out the availability of resistantseedling stock since greater than 95%of seedlings can be killed on sites ofhigh incidence of blister rust. In therust resistance programs, trees wereselected across an array of land owner-ship, and seedling offspring of theseparent trees were inoculated (infected)with the rust pathogen to identify par-ents and families with resistance. Seedorchards of resistant trees have beenestablished by various groups includ-ing the USFS and Bureau of LandManagement (BLM), as well as state,tribal, and private organizations.Recent data suggest the BLM Horningseed orchard, composed primarily ofparents from the western Cascadesarea of northern Oregon and southernWashington, produces seedlings withthe highest level of rust resistance inwestern white pine, suitable for plant-ing in many areas of western Oregon

Saving Our Trees and Foreststhrough Resistance Breeding

S

PHOTO COURTESY OF EVAN HECK

Controlled pollinations with western white pine at Washington StateDepartment of Natural Resources’ Meridian seed orchard. The full-sib seedproduced in these pollinations will be used in blister rust resistance tests toprovide advance-generation selections for rust resistance.

and Washington. In the RockyMountain Region, where westernwhite pine forests once dominatedmoist, mid-elevation sites, landownershave been planting resistant seedlingsfor decades. Many of these stands arewell stocked and are approaching har-vest age, while others have succumbedto intense rust pressure or failed forother reasons. Additional cycles ofresistance breeding are underway tofurther increase the level of resistancein the orchard seed available to forestmanagers.

In addition to breeding for rustresistance, the programs strive tomaintain both genetic diversity andadaptability within the species, andthe geographic range of each species isdivided into breeding zones to helpfacilitate this. The product for refor-estation is thus generally a geneticallydiverse mix of seed from orchardsusing many parent trees from thebreeding zone to be planted. Field tri-als of the resistant seedlots play animportant role. One notable series offield trials of western white pine seed-lots established by the WashingtonState Department of NaturalResources and other partners andcooperators examines blister rustresistance as well as well as the adapt-ability of seedlots from different geo-graphic areas on sites from Oregon toBritish Columbia. In addition, the fieldtrials serve as sentinel plantings tomonitor for other damaging agentsand as demonstration plantings forconservation education.

Programs to develop blister rustresistance in whitebark pine startedmore recently, but are making greatstrides. Whitebark pine, a high-eleva-tion conifer in western North America,is proposed for listing under theEndangered Species Act, with a deci-sion expected in 2019. The identifica-tion of resistant parent trees anddevelopment of blister rust resistantpopulations would greatly facilitatesuccess of restoration of the species.Restoration plantings of whitebarkpine using rust resistant seedlingshave been started on USFS andNational Park lands, and genetic trialsand conservation plantings on BLM,WA DNR and British ColumbiaMinistry of Forests, Lands and NaturalResource Operations lands. More lim-

ited resistance screening of the otherfive western species of white pines isunderway.

Spruce is an important species forreforestation. The program to developresistance to the white pine weevil isbased in British Columbia. Seed is nowavailable from orchards for both Sitkaspruce and interior spruce, and anincreased use of these species forreforestation has occurred. For interiorspruce, over 40 million seedlings fromorchard seed are being planted annu-ally in Canada with an estimate ofapproximately 30% less weevil damagerelative to unimproved (wild standseedlots) stock in the areas of highestincidence of the weevil. Increased lev-els of resistance are anticipated as theorchard is rogued and new resistantparents are added.

Port-Orford-cedar is a conifernative to northern California andsouthwestern Oregon but has beengrown horticulturally (often referred toas Lawson’s cypress) in many placesaround the world. The root diseasethreatens not only the native stands ofPort-Orford-cedar but also many ofthe horticultural plantings in bothNorth America and Europe. The pro-gram to develop genetic resistance,based at Dorena Genetic ResourceCenter, is a joint endeavor of the USFSand BLM with technical assistancefrom Oregon State University and withcooperation from many land man-agers. The program is one of thefastest developing applied forest dis-

ease resistance programs in the world,and significant increases in resistanceare available. The geographic range ofthe species has been divided into 13breeding zones, and innovative con-tainerized seed orchards are now pro-ducing resistant seed for many ofthese zones. Resistant seed is beingused for reforestation and restorationon USFS, BLM, National Park Service,state, county, tribal, and private lands.Breeding is underway to increase thelevel of resistance further.

Resistance is not immunity. In mostcases, only a percentage of seedlingsfrom orchards will be resistant, and itvaries by species. For resistance, suc-cess is a journey. The programs in thePacific Northwest are world leaders indevelopment of populations of treeswith genetic resistance to diseases orinsects. With a true integration ofresearch, applied tree improvement,smart reforestation and restoration,and support of the public, the road tosuccessfully maintaining these speciesin our forests continues and illumi-nates the pathway to follow as newinvaders affect our forests. The futureis sometimes cloudy, but resistance isan important tool in the path to futurehealthy forests. ◆

Richard Sniezko, an SAF member, is ageneticist for the US Forest ServiceDorena Genetic Resource Center inCottage Grove, Ore. He can be reachedat 541-767-5716 or [email protected].

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 29

PHOTO COURTESY OF RICHARD SNIEZKO

Breeding for resistance in Port-Orford-cedar containerized orchards.

BY BRIAN KLEINHENZ

hen I firststarted work-

ing in Alaska, I readan amazing accountof a timber cruiser inthe late 1930s. Thisforester was droppedoff on the north endof a remote island in early June. Hewas given a rough map, a few weeks’worth of canned food, and the prom-ise of resupply twice in that summervia air drop. This pioneer walked theberth and width of that island all sea-son collecting timber inventory dataalong the way. He spent all summer incontact with not a single soul, andafter being picked up on the beach atthe far end of the island in the fall,spent all winter working up his data. Inthis digital age, few of us can evenimagine being left alone long enoughto do all this. Where were his dead-lines? Why didn’t the world stop turn-ing when he missed his daily backupto the cloud? Many foresters relish ouralone time and cherish the peace ofthe woods. However, we are very luckyto have comradery and companion-ship with our fellow professionals in away impossible to that early Alaskantimber cruiser.

Foresters throughout the countryare more connected than ever. Weoften work on regional and nationalscales, and changing locations severaltimes during a career is commonplacein both the public and private sectors.The Northwest Office of the Society ofAmerican Foresters is an excellentexample of how our profession hasembraced this new reality of connect-

edness and used that energy to enrichthe traditional chapter, state, andnational levels of SAF organization.

In the Pacific Northwest, our fourstate societies have banded together toform a joint administrative structure inthe form of the Northwest Office(NWO). In a climate of stagnant mem-bership, it has been very useful for thesocieties to pool resources and shareservices. The NWO has two primarydeliverables. The first is the WesternForester, our very own regional publi-cation that covers issues relevant toforestry professionals in the PacificNorthwest. This has been a hugelypopular publication both with mem-bers and advertisers. Second, the NWOprovides a common source for admin-istrative services. The NWO maintainsan active website, interfaces withNational SAF, hosts conferences, servesas a stable point of contact for ques-tions from members and the public,and facilitates communicationbetween the various levels of SAF’sorganizational structure.

Together the Oregon, WashingtonState, Inland Empire, and AlaskaSocieties can support a stable, profes-sional staff to provide continuity inknowledge and services that a volun-teer base struggles with. With a modestannual budget of around $80,000, theNWO delivers excellent service andmaintains an office at the WorldForestry Center in Portland, Ore.Member assessments along with rev-enue generated by NWO publicationsand activities make this possible. A17-member committee with represen-tatives from all four societies guidesthe activities of the Northwest Office.This committee leverages a broad

range of experi-ence to coordinateefforts on thingslike policy actionand fundraising ina very effective manner. One of themost progressive efforts is the annualleadership conference organized bystate societies with support from theNWO. We collectively recognize theimportance of nurturing and support-ing the next generation of leaderswithin the forestry profession and ded-icate special annual training for thispurpose.

As a member of the Alaska Societyand Juneau Chapter, I have found thatparticipation in the Northwest Officehas enriched and expanded my experi-ence with SAF. While regular chaptermeetings are still the bedrock of myinterface to SAF, exposure to the NWOhas provided opportunities for collab-oration and leadership at a largerscale. It creates a much tighter knitand more effective Society. In manyways, the NWO has served to bridgethe gap between local reality andnational context. Our actions, positionpapers, fundraisers, and events are allmuch better with the input of regionaland national partners.

A century ago our vocation wascharacterized by individuals workingalone in the forest. Now we are collab-orators and innovators finding ways towork across geography. We can beproud that our Society is keeping pacewith these exciting times. I encourageeach of you to support our very ownNorthwest Office and get to know thenetwork of energetic individuals thatshare your passion for forestry. Yourdues and energy make this all possible.By reading the Western Forester andengaging in regional events you willsupercharge your career and forgesome valuable connections that willenhance your personal and profes-sional life. ◆

Brian Kleinhenz is past chair of theAlaska SAF. He can be reached [email protected].

30 WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017

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❑ Professional Forest Management ❑ Appraisals

❑ Timber Inventories and Cruising ❑ Mapping and GIS

Editor’s Note: To keep SAF membersinformed of state society policy activities,Policy Scoreboard is a regular feature in theWestern Forester. The intent is to provide abrief explanation of the policy activity—youare encouraged to follow up with the listedcontact person for detailed information.

SAF Hosts Riparian Science andManagement Workshop forPolicy Makers. Spurred by the trendof increasingly restrictive policies forriparian areas in the Pacific Northwest,a group of SAF and other naturalresource leaders organized a regionalworkshop on state-of-the-art riparianscience and management focused onpolicy makers and their staff. The all-day workshop was held on August 10 inPortland at the World Forestry Centerfor an audience of 105 that included anotable number of local, state, and fed-eral officials and staff. The program wasfully sponsored by SAF and severalother groups so that participants couldattend at no cost. A $3,000 Foresters’Fund grant was awarded in support ofthe event. Technical experts reviewedrecent study findings, and landownersand managers provided first-handaccounts that highlighted many com-plexities and challenges in managingthese unique areas in the broader land-scape. These observations helped showhow simplistic or overly restrictiveapproaches and policies for managingriparian areas can lead to missedopportunities for improving watershedconditions and resources, as well assome significant unintended conse-quences for both people and resources.Contact: Paul Adams, OSAF Policy chair,[email protected].

OSAF to Review PesticidesPosition Statement this Fall. Withthe coming expiration of OSAF’s posi-tion statement “Using Pesticides onForest Lands” (Dec. 2017), the state-ment will be reviewed for potentialchanges and updates to the discussionand supporting references. The use ofpesticides on forestlands remains a veryimportant issue that continues to stirpublic controversy and concern, partic-

ularly aerial spraying. For example,although its legality remains in question,a ballot initiative that bans aerial spray-ing was narrowly approved this spring byvoters in Lincoln County, Ore. All OSAFmembers are invited to review the exist-ing statement (www.oregon.forestry.org/oregon/policy/general) and passalong any comments to your local chap-ter officers or the Policy Committee.Contact: Paul Adams, OSAF Policy chair,[email protected].

New Western Oregon RiparianRules Now Effective, OtherRegions to be Considered. Withthe Oregon Board of Forestry’s (BOF)decision to increase restrictions inriparian areas on private forestlands inwestern Oregon, the Oregon Dept. ofForestry (ODF) finalized the specificrule language and related guidelines,which became effective on July 1.Because of persistent concerns aboutthe proposed rules and related researchand science concepts, OSAF submittedwritten input during the public com-ment period that preceded the finalrulemaking. Among the key messages:the added restrictions and complexityof the new rules are likely to discouragerather than encourage resource improve-ments through active management bylandowners and managers. In addition,the new rules rely heavily on tree basalarea as a surrogate for stream shade anddo not provide explicit allowance and

methods for shade-centric managementalternatives, which reflects limited con-sideration of more cost-effective policy tomaintain cool stream temperatures. Suchconcerns generally persist with the newrules and as potentially similar rules areconsidered by the BOF elsewhere inOregon. Details about the new westernOregon rules can be found at www.ore-gon.gov/ODF/Pages/LawsRules.aspxand the ODF website should provideupdates on rules considered for otherregions. Contact: Paul Adams, OSAFPolicy chair, [email protected].

OSAF Updates CommercialTimber Harvest Statement andPosition Statement Booklet.Earlier this year the OSAF ExecutiveCommittee approved an updated ver-sion of the position statement onCommercial Timber Harvest on PublicLands in Oregon. The revised textincludes more recent facts and figuresabout economic benefits of timber har-vest as well as some fine-tuning of theoverall text. The commercial harvestissue remains very timely given wide-spread management needs and costson federal forestlands, as well as thestatutory and long-held economic obli-gations to communities with large areasof nearby state or federal forests.Contact: Paul Adams, OSAF Policy chair,[email protected]. ◆

WESTERN FORESTER ◆ SEPTEMBER/OCTOBER 2017 31

Policy Scoreboard

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