Keynote Address – Brian J. Stanton

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Closing Keynote – Brian J. Stanton

Plant domestication in a changing world: Poplar markets, plantations, and science

OPENING

Good afternoon. I am Brian Stanton, GreenWood Resources’ Chief Science Officer. I

am here to close IPS-VI with a retrospective about how far we’ve come in developing

poplar plantations, markets, and science along with an outlook to the future, where we

are headed in poplar domestication and how we might arrive there. My talk will focus on

North America and I will not dwell too long on willow. But I will talk about what I know

best, poplar, and hopefully the information that I share today will be universally

applicable for this impressive international symposium.

Before I get started, I want to acknowledge the Faculty of Forestry at the University of

British Columbia, the IPS-VI conference organizing committee, and the officers of

IUFRO’s working group 2.08.04, Francisco Zamudio, Ron Zalesney, Deqiang Zhang,

and Theo Verwijst. We should commend them all and congratulate them for their warm

hospitality and for putting together such a terrific symposium. It has been a delightful

week. I think we can attest that the IPS movement is alive and well!

2014 is actually a well-chosen time to reflect on how far we’ve come in poplar and

willow domestication as it falls exactly 100 years after Augustine Henry’s first published

account of the successful controlled hybridization in the genus Populus at Kew

Botanical Gardens.i Coincidentally, 2014 also marks the 100th anniversary of the first

artificial forest regeneration program in North America, one that featured poplar. True

enough! Between 1894 and 1914 the Willamette Pulp and Paper Company established

400 hectares of black cottonwood (P. trichocarpa) plantations in the Pacific Northwest.ii

You can read about it in a 1952 issue of American Forests. Today, there are about 23

million hectares of forest plantations in the United States, principally loblolly pine in the

southeast and Douglas-fir in the Pacific coast region.iii And to think that it all started

with 400 hectares of black cottonwood. Two centennials, one in science and one in

industry, are to be celebrated. So it really is an appropriate time to take stock of the

past and to consider future possibilities.

Domestication is an applied process.iv It is the application of the science of genetics to

plant breeding. It is the manipulation of genetic variation from genomes to populations

in breeding and selecting varieties for commercial use. The theme of my talk this

afternoon is that markets drive industry that in turn drives science, which in its own way

drives more industry that sustains markets. This becomes an iterative process that

advances plant domestication. An example of this as seen in the history of the North

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America poplar plantation industry is a good place to start our search for the future of

poplar domestication.

Before we do, though, we ought to recognize that a lot is riding on this search! And

thankfully we’ve been given several guideposts that map our way to the future. In 2001,

Poplar Culture in North America was published, with a concluding chapter by Toby

Bradshaw and Steve Strauss that outlined a breeding strategy for the 21st century, now

already into its fourteenth year.v According to Toby and Steve, biotechnology ‒ both

genomics and transgenesis ‒ will lead the way in future poplar domestication. They

describe a coming global gene revolution. Continuing this message, Brian Ellis, Stefan

Jansson, Steve Strauss, and Gerry Tuskan told us in the 2010 text, Genetics and

Genomics of Populus, that genomics is developing so rapidly that the present state of

Populus domestication will look primitive in the year 2020.vi That is just six years away!

But tellingly, these same four authors conceded that we really can’t know how molecular

technology will be put into practice, because the application of science to poplar

domestication is at the mercy of social and economic factors. But in 2004, Reini

Stettler in an address to the First International Conference on the Future of Poplar

Culture told us that it’s not all about genes; in his judgment our community’s large pool

of scientific talent will be decisive in creating the expansive possibilities yet to come for

poplar.vii

Is the application of our science that dependent on the vagaries of society and the

marketplace? Is the transfer of technology really that serendipitous? That one

perceptive statement of Brian and his colleagues may be telling us what is needed to

push forward in poplar domestication: The application of science to plant domestication

will occur when future commercial ventures find themselves in need of, and willing to

pay for, the transfer of valuable technology. As quoted by Denis Murphy in his 2007

text Plant Breeding and Biotechnology: Societal Context and the Future of Agriculture,

the British scientist William Bate Hardy wrote in the early 20th century, “…applied

science is just as interesting as pure science but, it’s damned more difficult.”viii This

maxim is no less true in the 21st century: Research in poplar is having a heyday,

advancing discovery at a dizzying pace. But the application of this fine work is not

keeping pace. Applied science and poplar domestication throughout North America has

been exasperating. Why is this?

In part, the answer lies in the history of the poplar-using industry. And over the last 20

years – about the time the IPS movement was initiated in 1995 – we have watched a

rapid rise and an equally swift decline in this industry in North America. The history is a

fascinating story that played out in three broad regions of the U. S. and Canada during

the latter part of the previous century and continues today. It is a tale with three lessons

that instruct us how we may yet achieve the future of poplar domestication.

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THE HISTORY OF POPLAR INDUSTRIAL DEVELOPMENT

Commercial development of the poplar-growing industry in North America first took hold

nearly 60 years ago in the lower Mississippi River Valley when several companies

elta atch, U.S. Gypsum, Crown ellerbach, and ackaging Corporation all started

planting eastern cottonwood (P. deltoides) in response to the disruption in the natural

cycle of flooding essential to the regeneration of cottonwood stands brought on by the

construction of levees and revetments.ix Between 1958 and 1961 the largest plantation

- Fitler Managed Forest – was begun by Crown Zellerbach near Vicksburg, Mississippi.

Later in 1974, Westvaco Corporation started planting cottonwood further north at the

confluence of the Ohio and Mississippi Rivers. Thirteen thousand hectares were

producing pulpwood by the early 2000s with new varieties released by the U. S. Forest

Service Stoneville, Mississippi Experimental Station and the Texas Forest Service

following a process of phenotypic selection and clonal testing.x These varieties ′ST66′

and ′S7C8′ for instance, increased yields by 20% relative to unselected materials.xi

The second region where the history of the poplar-growing industry played out was the

Pacific Northwest when Scott Paper began planting in the lower Fraser River Valley as

early as 1950 using several P. ×canadensis selections from European breeding

programs including, ′I-214 ′ and ′Robusta,′ and ′OP-41′ bred by the Oxford Paper-New

York Botanical Garden hybridization program. Then, in the latter part of the 1970s and

continuing throughout the 1980s, Reini Stettler began hybridizing phenotypic selections

of the endemic P. trichocarpa with clonal selections of P. deltoides and Japanese poplar

(P. maximowiczii) at the University of Washington that led to the release of such notable

varieties as P. ×generosa '49-177' and P. trichocarpa × P. maximowiczii ′282-183.′ By

the mid-1980s, Scott Paper had shifted its program to the Stettler P. ×generosa and P.

trichocarpa × P. maximowiczii selections, and James River Corporation tapped into the

same mix of varieties in building their 4,500-hectare plantation in the lower Columbia

River Valley near Clatskanie, Oregon. These varieties were phenomenally successful,

showing 50% increases in wood volume compared to local P. trichocarpa selections.

By the late-1990s, Kruger Products had inherited the Scott Paper program that had

since grown to 3,500 hectares xii, MacMillan Bloedel had added 3,200 hectares in

northwest Washington and British Columbia using the Stettler hybrids, and Boise

Cascade and Potlatch Corporations had established a combined total of 15,200

irrigated plantation hectares in the mid-Columbia River basin near Boardman, Oregon,

and Wallula, Washington, using an assortment of Stettler’s P. ×generosa varieties and

the Oxford Paper P. ×canadensis varieties.

The last region where the history of the poplar-growing industry unfolded was a broad

expanse of the U.S. upper idwest and Canada’s prairie and eastern regions.

Beginning in 1994 in the U. S., Champion and later International Paper and finally Verso

Paper developed a hybrid poplar program using European P. ×canadensis and P. nigra

× P. maximowiczii varieties ′DN2,′ ′DN5, ′ ′I 45/51,′ and ′NM6′ to support their pulp

and paper operations in central Minnesota.xiii This quickly grew to 10,000 hectares

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supported by the ongoing varietal development work of the U. S. Forest Service

Rhinelander, Wisconsin Experimental Station and the University of innesota’s Natural

Resources Research Institute (NRRI). Top NRRI selections such as P. ×canadensis

′99059016′ now exhibit superior disease resistance with yield improvements of 70%

relative to the standard P. nigra × P. maximowiczii ′NM6.′

In Canada, Alberta-Pacific Forest Industries developed a 10,000 hectare poplar

plantation to support pulp-making operations at the Athabasca, Alberta mill. A

company-sponsored breeding program developed new hybrid varieties of both poplar

and aspen to improve winter hardiness, drought tolerance, growth rate, Septoria

resistance, and wood density. Further east, the Quebec provincial government has

funded the work of the Directorate of Forestry Research in hybrid improvement since

1969.xiv Improved varieties of multiple taxa including the P. deltoides × P. maximowiczii

taxon are used today across 6,000 industrial hectares managed by Domtar and

Norampac for pulp and paper and containerboard.xv In the Canadian prairie region,

Agriculture and Agri-Food’s Agroforestry evelopment Center at Indian Head,

Saskatchewan, has a history of over 60 years of poplar domestication.xvi Breeding and

selection has been targeted primarily for shelterbelt and farm feedlot plantings

throughout the Canadian prairie. Today, several varieties ‒ OPW-177H-86 and OPW-

120H-86 ‒ originating in a P. deltoides × P. ×petrowskyana three-way taxon have been

selected for widespread use.

At the turn of the new century, approximately 65,500 hectares of poplar plantations had

been established in North America. Quite the accomplishment in 42 years. What led

these 11 paper companies to turn to poplar for their raw materials? While the reasons

vary, it is fair to say that in most cases the primary decision to invest in poplar

plantations was the desire to secure fiber supply at a time when traditional wood

resources appeared to be dwindling.xvii There were other reason of course reduced

bleach re uirements, superior fiber properties for fine paper grades, etc. but the first

impulse had been raw material security.xviii

There are several things that we should note in this history. First, each of these

programs invested heavily in applied tree improvement in one way or another. Nearly

all companies employed poplar geneticists with Ph. D. or M. S. degrees to breed and

select improved varieties. Nearly all allied with government or university labs to initiate

their programs. All considered their deployment varieties confidential intellectual

properties, but none ever developed a commercialization strategy to capitalize on this

IP. And most, at one time or another, belonged to a university-led cooperative that was

promoting the development of genomic tools or genetic transformation technologies.

Clearly, there is a lesson we can draw here: The poplar-growing industry placed an

astonishing priority on applied science to secure their paper businesses using the

classical approach to plant hybridization and selection combined with the

biotechnological approaches that were coming into vogue in the 1980s.

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And then, beginning in the late 1990s, it all began to fall apart. There were a multitude

of reasons for the industry’s demise, but the two most consistent ones appear to have

been: (1) the anticipated shortage in traditional fiber sources never fully materialized,

and (2) an increase in global competition in the communication paper sector. To remain

competitive, paper companies began to shed their poplar holdings. The first to go was

the MacMillan Bloedel program that initially spun off its plantations in 1998 into a

separate entity, Pacifica Poplars, later liquidated. Next to go was the Fitler operation

that was sold to a real estate developer in 1999. In 2000, James River was acquired by

Georgia Pacific, which then divested itself of the Clatskanie tree farm. In 2003 and

2004, Boise Cascade and Westvaco corporations were sold to private investment firms,

precipitating the sale of all or a substantial share of their poplar holdings.xix In 2007,

Potlatch Corporation converted to a REIT and gave up its poplar estate, considering it

no longer strategic. Finally, in 2012 and 2013, the last examples of the fully integrated

plantation-pulp mill model came to an end as Verso Paper sold its Minnesota properties,

Alberta-Pacific Industries discontinued its planting program, and Kruger Products began

selling off their Fraser Valley lands.xx Today, Boise Cascade’s scaled-down plantation

operation is the lone vestige of the paper industry’s poplar experience. The only

research and development programs to remain untouched by the demise were the

publically-funded ones in Saskatchewan and Quebec.

A breathtaking, albeit disappointing history! But one in which we uncover a second

lesson: An industry that adopted plantation operations most often as a strategic

response to constraints in customary sources of raw materials quickly gave up those

operations when supply and pricing forecasts for their traditional base of raw materials

returned to a new normal.xxi The poplar-growing operations were at risk if viewed

principally as a cost center.xxii

The back story here is that the forest products industry was undergoing a structural

change during this period in which companies were separating their timberlands from

their manufacturing base.xxiii The reasons have been described as: 1) the understated

market value of timberland holdings, 2) the need for investment capital for mill

improvements, and 3) the increased supply of wood from non-company lands.xxiv The

institutional investment community soon took note that timberland might be a profitable

new asset class for several reasons.xxv First, timberland investments diversified

financial-asset portfolios. Second, they provided inflation protection simply from the fact

that wood volume accumulates with each growing season even when prices are

increasing in the same economy. Third, initial rates of return exceeded the Security

Market Line, approximating returns of 12 to 14% per year. xxvi By the mid-1990s,

institutional investors held more than $2.5 billion of market value in traditional U.S.

timberlands, consisting primarily of coniferous forest holdings. Eventually, return rates

fell and investors began to look elsewhere for new opportunities in emerging timber

markets. This created an opportunity for an inventive investment product that set the

stage for hybrid poplar’s reemergence into two new markets.

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About this time, the sawn wood and panel markets were seeking alternatives to alder,

aspen, and basswood for appearance grade dimensional hardwood lumber and a

replacement for yellow poplar in producing decorative hardwood veneer cores. Both

markets settled on hybrid poplar when GreenWood Resources created the GreenWood

Tree Farm Fund (GTFF). Working with institutional investors in 2008, GTFF acquired

17,000 hectares of the former poplar plantations of three pulp and paper companies in

Oregon and Washington, converted the silviculture to the production of quality saw- and

peeler logs, constructed a state-of-the art sawmill, and enlisted Collins Company as a

GTFF investor to manage the annual production of 80 million board feet of kiln dried

lumber. A veneer mill was added in 2013 under a separate arrangement with Columbia

Forest Products to annually produce 45,000 cubic meters of veneer. Now a fourth

market – renewable energy feedstock – awaits.

What is not to be missed here is poplar’s uni ue combination of traits that allow

adaptability to such a wide range of products. It is hard to image any other North

American species matching poplar’s tremendous juvenile growth rate, low cost

vegetative propagation, coppice regeneration, relatively low lignin content, and light

weight, straight grain, fine textured, bright wood that promote it in all four markets. The

investment community also recognized this. But before committing to an equity position

in GTFF, investors who were increasingly aware of the genetic potential of varietal

forestry,xxvii questioned whether their investment would be underwritten by a

hybridization program to increase yield and wood quality, and maintain pest resistance.

The answer of course was “yes.” For in its structuring of GTFF, GreenWood

consolidated many of the U. S. industrial genetic improvement programs from the pulp

and paper era. These included breeding populations of three hybridization species,

base populations and production varieties of multiple intra- and inter-specific taxa,

greenhouses, research equipment, data files, and personnel with critical know-how.

GTFF investors’ enlightened opinion of genetics was refreshing, validating the worth of

the intellectual property that had been developed by the paper industry. This illustrates

a third lesson: The market will respond to creative investment vehicles that provide a

compelling case of financial performance, market appeal, and, most revealingly, the

clout of technology transfer.

TECHNOLOGY TRANSFER

How did GTFF investors come to appreciate the consequentiality of technology transfer

in poplar? I’d like to think that they studied their history, perhaps the history of applied

plant breeding and its success in transforming American agriculture and everyday

American life during the 20th century in the form of a hybrid plant: hybrid corn!

Hybrid corn; what a story, what a stupendous example of effective technology transfer!

As early as 1881, W. J. Beale first demonstrated the potential of inter-varietal F1 corn

hybrids. Between 1917 and 1923, the first commercial hybrid varieties were developed

and released based upon the work of geneticists George Shull and Edward East.xxviii

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Wholesale use of hybrids quickly followed, and by 1930 appreciable yield increases

were being recorded.xxix By 1953, hybrid varieties accounted for 95% of the corn crop

grown in the Midwest As a result, yield increased throughout the 20th century from

1,600 kilograms per hectare to 9,400 kilograms per hectare, a nearly six-fold upturn.xxx

The effect on everyday life was stunning: Consumer real prices fell by one-half, and

disposable income spent on food products fell by two-thirds during this time.xxxi It

should be inarguable that the investment in applied plant breeding was a socially

profitable one brought about by the public sector: The U. S. Congress established the

Land Grant University system with the Morrill Act in 1862 that created agriculture as a

scientific profession, and the State Agricultural Experiment Stations were created in

1887 to test the new F1 crop varieties and disseminate seed to local farmers.xxxii It is a

great story: Public investment, good science, effective technology transfer, agriculture

remade. What about poplar? Is this same dramatic effect possible? I know it is, but

there is a sense of urgency in bringing it about. Let me explain.

The renewable transportation fuels industry is poised to take wing in the U. S. using

public investment: The U. S. Department of Agriculture-NIFA is supporting seven large

regional energy research and development projects across the country to bring this

about. Three include poplar or willow as featured feedstock crops. Poplar technology

transfer will be a big part of realizing this industry’s future, for among other reasons

feedstock production accounts for fully a third of the total operating cost of a large bio-

refinery.xxxiii But as we anticipate the renewable biomass market and public investment

in poplar and willow, we would be well advised to recall the past, for there are parallels

that should give us pause: The pulp and paper industry embraced poplar as an

incremental fiber supply to forestall a shortfall in an accustomed raw material base.

When the shortage failed to materialize, poplar lost its attractiveness as a replacement

fiber supply. Sounds a lot like the energy industry: A global commodity market

returning to its traditional resource base, in this case shale oil and gas reserves

revitalized by a mid-20th century technology, hydraulic fracturing. So what will happen

next? Is the promise of poplar in the renewable energy market likely to go

unfulfilled?xxxiv I don’t think this will be the case. The answer lies in how we fund our

science and husband the transfer of its technology. And that is why this conference is

so vital to poplar’s future domestication possibilities.

TOWARDS A MORE COMPREHENSIVE DOMESTICATION PROGRAM

So if the next stage of poplar domestication is contingent on an effective transfer of

technology that embodies all of the wonderful science reported here this week, what

needs to be done? One tactic in charting a path to the future would have us apply past

lessons to tomorrow’s challenges.

Challenge: Rapid changes in domestication priorities will require poplar breeders to

effectively integrate more sophisticated, biotechnological methods to accelerate genetic

improvement if they are to survive the vagaries of industrial markets. Genetically

improved varieties can guarantee success in each of the five markets in which poplar is

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used (i. e. communication grade papers, appearance grade lumber, hardwood core

veneer, oriented strand board, energy feedstock). We are witnessing the rapid

development of molecular breeding tools predicted by Brian Ellis and his co-authors. It

is quite certain that we will soon have the capability to score millions of markers across

individual genomes at a reasonable cost, resulting in haplotypes characterized for large

numbers of loci that explain a substantial portion of phenotypic variation. Molecular-

based selection could soon be a reality for poplar, leading to a significant reduction in

test rotations. Research in transgenic poplar is moving forward: The Department of

Agriculture’s A HIS’ Biotechnology Regulatory Services’ website lists a large number of

permits for field tests of transgenic poplars for traits as diverse as growth, flowering,

drought tolerance, wood quality, light response and more. In Europe, Swetree has

approximately 40 gene constructs in field tests featuring growth, water use efficiency

and more. Futuragene is testing genes in poplar for yield improvement and insect

resistance.

This is truly thrilling stuff. But the path forward is likely to be fraught with complications

that challenge good decision-making in the design of future domestication strategies.

Let me cite two papers from the forest genetics literature that together illustrate how

challenging the integration of molecular tools into conventional breeding programs may

be. One paper describes the domestication of forest trees just prior to the advent of

molecular tools, the second well into the advance of molecular tools. The first is Bill

Libby’s paper, “ omestication Strategies for Forest Trees” published in 1973 in the

Canadian Journal of Forest Research that explicated a variety of domestication

pathways topical for the times, from phenotypic selection and controlled breeding

through progeny testing, parental selection, seedling and clonal seed orchard

establishment, and plantation deployment.xxxv Forty years later in 2012, Antoine

Harfouche and his colleagues published “Accelerating the omestication of Forest

Trees in a Changing World” in Trends in Plant Science that nicely enumerated all of the

wonderful molecular technologies under development.xxxvi The message they deliver

would have been unrecognizable in 1973: The optimum domestication approach

integrates transgenesis and genomic selection, the latter using next generation

sequencing to scan a sufficiently large number of markers at low cost to circumvent

linkage disequilibrium. Both are well written papers but their construction is interesting.

Whereas the 1973 domestication paper has a single author, Bill Libby, it does not cite a

single reference for the reader’s supplemental research, and includes just one table and

one flowchart – neither footnoted – to illustrate his arguments. Harfouche’s 2012 paper

includes six co-authors, 63 references, a glossary defining the new technologies, and

two sidebars detailing the technology transfer process. Let’s all agree that

domestication has gotten a whole lot more complex over the 40-year timescale in which

these two papers appeared!

We should recognize from the 2012 paper that genomic selection, association genetics,

and transgenesis will surely play a pivotal role in poplar domestication.xxxvii But, they will

supplement and not replace traditional breeding and selection. The cornerstone of

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poplar breeding will always firmly rely on controlled hybridization, field-based selection,

and quantitative genetic analyses.xxxviii Investments into molecular techniques will be

worthwhile only if there is a parallel investment in developing the hybridization and

selection programs to which these marvelous tools can be applied. What concerns me

is not whether we will perfect the new technologies. (I believe there is no question

here.) What concerns me, rather, is the absence of any effort to bring together at a

national level, the full spectrum of genetic resources that are manipulated and

domesticated with conventional and new technologies that lead to the transfer of

technology that drives industry and opens markets. Without this, the investment into

genomics and transformation sciences will not pay off.

Challenge: A collaborative approach to poplar domestication should yield community

benefits that amplify the individual research and development efforts of the government,

academic, and industrial sectors. I can only imagine how much further along we would

be in the U. S. national poplar domestication program had the paper industry adopted a

collaborative improvement approach during the 1980s 1990s. Perhaps we would

now see more extensive provenance sampling of our key hybridization species, greater

advancement in reciprocal recurrent breeding of parental species, more broadly-based

field trials of elite varietals, and greater exploratory hybridization of novel taxa to widen

the genus’ adaptability.

Can this still be accomplished? And how? One appealing thought is a coordinated,

publicly-funded national poplar improvement center. There is something of a precedent

here: Over the past several years, the U. S. Department of Energy’s SunGrant

Partnership program has supported poplar varietal development under the leadership of

Tim Rials, University of Tennessee, and Bill Berguson, University of innesota’s

Natural Resource Research Institute.xxxix The priority has been the traditional route to

poplar improvement for the renewable biomass industry. This program might be the

logical choice to expand to a new mission to develop a national poplar domestication

center. Yet we cannot expect the SunGrant Partnership program to assume a

sustainable funding role, because its mission is to provide for the information needs of

the Department of Energy rather than the funding of a long-term project such as the one

I am proposing.

Perhaps then, a national domestication center could follow the university-industry

cooperative model that has worked so well for the southeastern pine industry.xl But

bringing this about in poplar will be a challenge: The North American poplar industry

pales in comparison with the southeastern pine industry in the strength of its economy,

the number of industrial players, the land area under management, the quantity of

manufacturing facilities, and so forth. But these notwithstanding, the advantages of a

cooperative are significant, especially where a strong community network is in place as

it is with poplar. But it will be hard to muster industrial support for a poplar cooperative

until a market pull for substantial quantities of poplar raw materials develops.xli That is

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today’s reality. I believe that such market pull will come, but what can be done until that

time? That brings us to the final challenge.

Challenge: Advancement in the science and technology of poplar genetics offers a

significant inducement for the addition of a novel funding source to develop a national

hybridization and varietal selection initiative. First, the good news: Public investment

in poplar genomics is robust. The epartment of Energy’s Bioenergy Science Center at

the Oak Ridge National Laboratory is funded at $25 million annually, an essential

component of which supports the work of Gerry Tuskan and Steve DiFazio and their

colleagues in poplar molecular science. Additionally, the Plant Feedstock Genomics

program, jointly funded by the Departments of Agriculture and Energy provides up to

$500,000 in annual funding for individual, multi-year projects. And here in Canada,

Genome Canada and Genome B.C. together have provided $10 million over a seven-

year period to support the poplar genomics work of Carl Douglas and Shawn Mansfield

at the University of British Columbia as it can be applied to the development of

Canada’s renewable bioenergy industry. These are wise public investments. In the

private sector, three companies ArborGen, Futuragene, and Swetree have

developed advanced research labs with poplar transformation capabilities. And a

fourth Rapid Genomics is offering genomics assisted breeding services in poplar.

Support from government and industry for the molecular realm is assured.

Contrast this, however, with public support for the traditional improvement route. We

can point to the provincial program in Quebec as a first-rate example of sustained public

investment in poplar hybridization and selection.xlii But in the United States there is no

comparison: It is only the aforementioned epartment of Energy’s SunGrant

Partnership program that lends supports to traditional hybridization. Even then, the

funding level is quite modest and it is uncertain whether the program will be extended

beyond 2015. And in the private sector only GreenWood Resources is pursuing both

poplar population improvement and elite population development on an ongoing basis in

service to the private e uity investors that participate in the company’s tree farm funds.

So if the publically-funded model and the industrial-university cooperative model will be

a challenge to bring about, and if the private sector effort falls short of the broader

commission of a national domestication program, what is the answer? It may be a new

trend called science philanthropy. Earlier this year The New York Times reported on a

profound change taking place in the way science is being funded throughout the U. S.xliii

Some of the very richest Americans are becoming science patrons to advance social

progress where they find a cause that appeals to their passion and their entrepreneurial

spirit. Examples include Microsoft co-founder Paul Allen, Eric Schmidt of Google,

Lawrence Ellison of Oracle, businessman and investor Ronald Perelman, financier

ichael ilken, and of course, Bill Gates. The lion’s share of the philanthropically

funded science is going to the search for cures for human illnesses. Even the American

Association for the Advancement of Science recognizes that the new philanthropists will

play a vital role in advancing translational science.xliv Can you imagine a group of

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philanthropists being inspired by and enthusiastic about a plant domestication program

that leads to a secure and sustainable renewable energy supply? One that also impacts

climate change? One that revitalizes our rural economies? Wouldn’t that be

something!

Do you think this idea whimsical or farfetched? It might be. But on the other hand it

may not be so unrealistic. We’ve seen this before with an equally compelling issue:

Human starvation. About 55 years ago, right about the same time that the first industrial

poplar plantations were being installed in the lower Mississippi River Valley, the

philanthropic sector took note of the desperate needs of the developing world and

began to fund advances in crop domestication that fueled the Green Revolution,

increasing global agriculture production perhaps saving over a billion people from

starvation by development high-yielding varieties of cereal grains.xlv The Revolution

really began in earnest in the 1960s when the Rockefeller Foundation formed an

alliance with the Mexican government to improve wheat genetics under the guidance of

Norman Borlaug. Later in the 1970s, the Ford Foundation worked with the Philippine

government to improve rice genetics. Like Borlaug the wheat breeder, rice breeding

had its champion in Henry Beachell. Results were spectacular: wheat and rice yields

trebled in a decade. In 1971, the Food and Agriculture Organization (FAO) and the

World Bank established the Consultative Group on International Agricultural Research

(CGIAR) as a global network for crop improvement. In 2014, the Gates Foundation and

the Carlos Slim Foundation provided a $25 million grant to CGIAR to keep agricultural

science moving forward in developing countries worldwide.xlvi The role of the

philanthropic investor continues in support of world food production. I suspect that the

philanthropist are now searching for new horizons.

CLOSING

So this is where domestications needs to go in the future: A nationwide center of plant

material development that features the classical approach of hybridization and selection

at its core, enhanced with molecular tools and support from stable funding mechanisms

provided through a combination of the public good, industry, private equity, and maybe

a new brand of philanthropy. And there is an exigency to achieve this.

The most recent International Poplar Commission (IPC) report shows that the worldwide

area of poplar and willow plantations increased from 4.8 to 5.3 million hectares between

2008 and 2012. Many IPC member nations believe that this area will increase further.

This is in line with FAO’s new publication, Poplars and Willows: Trees for Society and

the Environment that forecasts greater reliance on the domestication of Populus genetic

resources. This is timely: Despite the existence of many fine breeding programs in

Asia, Europe and North America, many timeworn varieties remain in commercial use.

My hope is that IPS-VI does not conclude with this address. Rather, I trust that the

organizers will poll the delegates to gauge our community’s interest in supporting a

national or regional or global center of poplar and willow domestication. Leadership may

Keynote Address – Brian J. Stanton

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originate in a variety of ways: IUFRO’s poplar and willow genetics working group, I C’s

working group on poplar genetics, a national poplar commission such as the one here in

Canada, a government agency, or some enterprising individual. The key is to begin a

dialogue and to take action. This will give IPS its relevance.

Thank you very much.

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ENDNOTES

i Henry, A. 1914. The artificial production of vigorous trees. Journal of the Department of Agriculture and Technical Institute, Irish Free State 15: 34-52. Henry’s report lists two successful inter-specific Populus crosses, P. angulata × P. trichocarpa and P. angulata × P. nigra var. betulifolia. P. angulata is now known as P. deltoides. Henry is the naming authority of P. angulata × P. trichocarpa, now known by its hybrid binomial, P. ×generosa. ii riaulx, A. W. 1952. Oregon’s planted cottonwoods. American Forests. 58: 10-11, 66-68. 70. iii Smith, W. B., Patrick, M.D., Perry, C. H., and Pugh, S. A., 2009. Forest Resources of the United States, 2007. Gen. Tech. Rep. WO-78. Washington, DC: U.S. Department of Agriculture, Forest Service, Washington Office. 336 p. iv Libby, W.J. 1973. Domestication strategies for forest trees. Canadian Journal of Forest Research 3, 265-276. v Bradshaw, H.D., Jr. and Strauss, S.H. 2001. Breeding strategies for the 21st century:

domestication of poplar. In: Dickmann, D.I., Isebrands, J.G., Eckenwalder, J.E. and J.

Richardson (eds) Poplar Culture in North America. National Research Council of Canada

Research Press, Ottawa, Ontario, Canada, pp. 383-394.

vi Ellis, B. Jansson, S., Strauss, S. H., and Tuskan, G. A. 2010. Why and how Populus became

a model tree. In Genetics and Genomics of Populus. Edited by S. Jansson, R. P. Bhalerao, and

A. T. Groover. Springer, New York, pp. 3-14.

vii Stettler, R. F. 2004. The actual and the possible – The role of science in poplar’s future. In: First International Conference on the Future of Poplar Culture. Italian National Poplar Commission (compilers) FAO, Rome, p. 18. viii Murphy, D. 2007. Plant breeding and biotechnology: Societal context and the future of agriculture. Cambridge University Press. Cambridge, U. K. 423 pp. William Bate Hardy, British biologist and food scientist, 1864 – 1934.

ix The Lower Mississippi River is a stretch of 1,600 kilometers between the confluence of the Ohio River and the Upper Mississippi River at Cairo, Illinois and the Gulf of Mexico. There are no locks or dams on the Lower Mississippi. The river is, however, constrained by levees to control flooding and secure a navigation channel for barges.

x Mohn, C. A., W. K. Randall, and J. S. McKnight. 1970. Fourteen cottonwood clones selected for midsouth timber production. USDA Forest Service Research Paper SO-62, 17 pp. xi Personal communication. Pat Weber, former manager of Fitler Managed Forest. xii Personal communication, Peter McAuliffe, former Scott Paper/Kruger Products forester. xiii Hansen, E. A., Ostry, M. E., Johnson, W. D., Tolsted, D. N., Netzer, D. A., Berguson, W. E. and Hall, R. B. 1994. Field performance of Populus in short-rotation intensive culture

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plantations in the north-central U. S. Res. Pap. NC-320. St. Paul, MN: U. S. Department of Agriculture, Forest Service, North Central Experiment Station. 13 p. xiv Vallee, G. 1969. A general program for forest tree improvement in Quebec. Tree improvement Branch, Quebec Ministry of Lands and Forests. 59 p. xv Perinet, P. 2007 The poplar breeding program in Quebec. In: Perinet, P., Perron, M. and

Belanger, P. (eds) Poplar Culture: A Collaborative Effort from Clone to Mill. 2007 Annual

Meeting of the Poplar Council of Canada. Ministère des Ressources naturelles et de la Faune

du Québec, Direction de la recherche forestière, Québec, Canada, pp. 11-12.

xvi Cram, W.H. 1960 Performance of seventeen poplar clones in south central Saskatchewan.

The Forestry Chronicle 36, 204-224.

xvii In many cases the curtailment of harvest operations from federal lands in the U. S. resulted in the closure of sawmills and a reduction in sawdust and shavings that ordinarily would have been available to the pulp and paper industry. In other cases, an imbalance in the age class distribution in alder and aspen created an impending shortfall in hardwood fiber that papermakers needed for the manufacture of white, uncoated, free sheet papers. xviii Poplar was used for a range of lightweight, coated and uncoated grades of specialty newsprint. The comparatively low wood density proved well suited to the thermo-mechanical process pulping process where energy could be conserved during chip refining. Additionally, poplar’s characteristic bright wood was preserved in mechanical pulps requiring minimal hydrogen peroxide bleaching. The chemical process was used in pulping poplar where the short and relatively wide, thin-walled fibers were suited to the manufacture of high-quality bond paper grades. (Chemical fibers are low in coarseness and collapse easily during sheet formation with longer softwood fibers resulting in a smooth, dense formation with few surface voids, superior opacity, and good print retention.) xix Westvaco was sold to Cerberus Capital Management and Boise was sold to Madison Dearborn Partners xx Boise Cascades’ Wallula, Washington pulp and paper mill maintains a scaled-down poplar plantation operation as the lone vestige of the paper industry’s involvement with poplar. xxi Although other factors figured into the fate of the paper industry’s poplar plantations, the supply balance of hardwood fiber from native stands seems to have been one of the main contributory factors. xxii A cost center is a business unit that does not generate direct profits but adds to the cost of running a company. xxiii Binkley, C. S., Aronow, M. E., Washburn, C. L. and New, D. 2005. Global perspectives on intensively managed plantations: implications for the Pacific Northwest. Journal of Forestry 103: 61-64. xxiv Binkley, C. S., Raper, C. F., and Washburn, C. L. 1996. Institutional ownership of US timberland. Journal of Forestry. 94: 21-28.

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xxv Institutional investors are organizations that place sizable investments into securities, timberlands, and other real assets. Typical investors include banks, insurance companies, university endowments, retirement or pension funds, hedge funds, investment advisors and mutual funds. Timber Investment Management Organizations (TIMOs) came about when Congress mandated broader diversification for institutional investment portfolios. At the end of 2009, TIMOs had about $24 billion invested in U.S. timberland, up from just $1 billion in 1989. (Spears, L. D. 2011. Timber Investing: The Inflation Hedge That Pays Off in Every Type of Market. Money Morning). The growth of institutional ownership of timberlands has been driven by changes to U.S. accounting rules that assume that trees not only don’t grow but rather depreciate over time. (Binkley, C. S., Raper, C. F., and Washburn, C. L. 1996. Institutional ownership of US timberland. Journal of Forestry. 94: 21-28.) xxvi “Since 1910, the value of timberlands as an investment has risen faster - and with less

volatility - than stocks as measured by the Standard & Poor's 500 Index. Since 1987 alone - in

spite of minor losses in 2010 due to the U.S. housing slump - the Timberland Index maintained

by the National Council of Real Estate Investment Fiduciaries has risen roughly 15% per year,

compared to an annualized return of just 9.61% for the S&P 500.” (Spears, L. D. 2011. Timber

Investing: The Inflation Hedge That Pays Off in Every Type of Market. Money Morning.

xxvii Merkle, S. and Cunningham, M. 2011. Southern hardwood varietal forestry: a new approach to short-rotation woody crops for biomass energy. Journal of Forestry 2011: 7-14. xxviii Murphy, D. 2007. Plant breeding and biotechnology: Societal context and the future of agriculture. Cambridge University Press. Cambridge, U. K. 423 pp. xxix Gibson, M. 2012. The feeding of nations: Redefining food security for the 21st century. CRC Press. 684 pp. xxx Troyer, A. F. 2006. Adaptedness and heterosis in corn and mule hybrids. Crop Science 46: 528-543. Troyer, A. F. 1998. Background of U. S. hybrid corn. Crop Science 39: 601-626 xxxi Gardner, B. L. 2002. American agriculture in the twentieth century. How it flourished and what it cost. Harvard University Press. 388. Pp xxxii Murphy, D. 2007. Plant breeding and biotechnology: Societal context and the future of agriculture. Cambridge University Press. Cambridge, U. K. 423 pp. xxxiii Gustafson, Rick. AHB Executive Team Strategic Planning Meeting. Presentation, Portland January 30, 2014. xxxiv A recent poll on alternative energy sources (Yale Project on Climate Change Communication Project, Climate Progress, Feb., 2014) reported that many are willing to pay more for the benefits of a low-carbon renewable energy industry and included the following statistics:

· 83 per cent of Americans want their country to make an effort to reduce global warming, even if it has economic costs. · 60 per cent of Americans think the US should act "regardless of what other countries do;" · 67 per cent want to regulate carbon dioxide as a pollutant; · 59 per cent want to cut fossil fuel subsidies entirely; · 60 per cent think cutting renewable subsidies is a bad idea;

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· 72 per cent want more research funding for renewable energy; · 59 per cent of Americans want the EPA to regulate carbon pollution from power plants; · Different versions of a cap-and-trade system or a carbon tax gained the support of almost half of respondents.

xxxv Libby, W. J. 1973. Domestication strategies for forest trees. Canadian Journal of Forest Research 3: 265-276. xxxvi Harfouche, A., Meilan, R., Kirst, M., Morgante, M., Boerjan, W., Sabatti, M., and Scarascia Mugnozza, G. 2012. Accelerating the domestication of forest trees in a changing world. Trends in Plant Science 17: 64-72. xxxvii The integration of new technologies will be challenging as, for example, in the choice

between genomic selection and association genetics. Right now genomic selection is in vogue.

But will it be the more efficient route in breeding for heterosis if first-generation inter-specific

performance is controlled by a modest number of traits? Shouldn’t association genetics work

as well in situations where we have a good grasp of inheritance and a good understanding of

the polymorphism directing trait expression? In a sequenced organism like poplar that is paired

with an exhaustive understanding of physiology, morphology, and pathology perhaps we won’t

require tens of thousands of SNP markers, maybe 100 will do using an association approach. If

the causative polymorphism are found, known, and constant, then an association approach

should work precluding the need to recalibrate selection models necessary with genomic

selection. On the other hand, biomass production and other commercial traits are surely under

polygenic control, and it is probably too difficult to tease out all of the polymorphisms with an

expectation of explaining a large portion of phenotypic variation using an association genetics

approach. Here, genomic selection may be the correct route, because of the inherent difficulty

in deducing positive and negative allele function. But we can’t forget that genomic selection

models are being built with no assurance that the genetic structure of today’s training

populations will match those of the breeding populations for which they are intended for the

projection of genomic estimated breeding values. And it is unknown how many generations we

will be able carry such models before predictive power diminishes. Thus I think it prudent to

keep both options in play so that whatever form the application of molecular technology takes,

our conventional tree improvement programs can anticipate the logistical hurdles ahead. The

integration challenges transformation science will face are regulatory in nature, reflecting not

only bona fide environmental concerns but also society’s biases. erhaps the best we can do

now to advance this branch of the new technologies is to advocate for the USDA to adjudicate

applications for the release of transformed poplar varietals on the basis of science, as opposed

to an agenda rooted in politics, philosophy, or emotion.

xxxviii Murphy, D. 2007. Plant breeding and biotechnology: Societal context and the future of agriculture. Cambridge University Press. Cambridge, U. K. 423 pp. van Buijtenen, J. P. 2001. Genomics and quantitative genetics. Canadian Journal of Forest Research. 31: 617-622. xxxix That component of the OE’s SunGrant artnership program that supports poplar varietal improvement is led by the University of innesota’s Natural Resource Research Institute. Other participating organizations are ArborGen, GreenWood Resources, Michigan State University, and Mississippi State University.

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xl Byram, T. B., Mullin, T. J., White, T. L. and van Buijtenen, J. P. 2005. The future of tree improvement in the southeastern United States: alternative visions for the next decade. Southern Journal of Applied Forestry. 29: 88-95. xli The annual operating budget of a university-industrial poplar genetic improvement cooperative is estimated to be $1.4 million to conduct a hybridization program at two locations, as well as an extensive field-testing program throughout the United States. This assume in-kind contributions provided by coop members. xlii Unlike the industrial poplar breeding programs in the U.S., the government-sponsored

program in Quebec has endured for nearly 55 years. Why is this? It can be partly explained by

the fact that management responsibility for crown lands that represent 89% of Quebec’s forest

land base, resides with the Ministry of Natural Resources. This entity considers tree

improvement the foundation of its reforestation programxlii. The Directorate of Forestry

Research has received, therefore, sustained support for a multi-species breeding program;

though poplar regeneration accounts for less than one million cuttings per year, the Directorate

has been able to conduct cost-effective poplar hybridization and selection since 1969 as one

facet of a broader improvement program. A surge in the consumption of aspen, up to 5.4 cubic

meters in 2004, by the plywood and OSB industries has further guaranteed ongoing support for

the Quebec poplar breeding program.

xliii Broad, W. J. 2014. Billionaires with big ideas are privatizing American science. The New York Times, March 15, 2014. xliv McNutt, M. 2014. The new patrons of research. Science 344: 9. xlv Murphy, D. 2007. Plant breeding and biotechnology: Societal context and the future of agriculture. Cambridge University Press. Cambridge, U. K. 423 pp. xlvi Global research network raises $1 Billion for its Centers. Science