Am erican Journal o f
American Journal of Botany 101(10): 1597-1600,2014.
C e le b r a t in g lO O y e a r s ' ^ 11 9 1 4 - 2 0 1 4
INTRODUCTION
S peaking of food: C onnecting basic and applied plant science1 B riana
L. G ross2, E lizabeth A.
K ellogg 3,
and
A llison
J. M iller4’5
departm ent of Biology, University of Minnesota Duluth, 207 Swenson Science Building, 1035 Kirby Drive, Duluth Minnesota 55812 USA; 3Donald Danforth Plant Science Center, 975 North Warson Road, St. Louis, Missouri 63132 USA; and departm ent of Biology, Saint Louis University, 3507 Laclede Avenue, St. Louis, Missouri 63103-2010 USA The Food and Agriculture Organization (FAO) predicts that food production must rise 70% over the next 40 years to meet the demands of a growing population that is expected to reach nine billion by the year 2050. Many facets of basic plant science promoted by the Botanical Society of America are important for agriculture; however, more explicit connections are needed to bridge the gap between basic and applied plant research. This special issue, Speaking of Food: Connecting Basic and Applied Plant Science, was conceived to showcase productive overlaps of basic and applied research to address the challenges posed by feeding billions of people and to stimulate more research, fresh connections, and new paradigms. Contributions to this special issue thus illustrate some interactive areas of study in plant science— historical and modem plant-human interaction, crop and weed origins and evolution, and the effects of natural and artificial selection on crops and their wild relatives. These papers provide examples of how research integrating the basic and applied aspects of plant science benefits the pursuit of knowledge and the translation of that knowledge into actions toward sustainable production of crops and conservation of diversity in a changing climate.
Key words: agriculture; agroecological research; applied science; basic science; botany; crop domestication; crop evolution; food production; Pasteur’s quadrant. The Food and Agriculture Organization of the United Na tions predicts that food production must rise by 70% over the next 40 yr to meet the demands of a growing population ex pected to reach nine billion by the year 2050 (UNFAO, 2009). The solution to this challenge will be multifaceted and certainly involve changes in what, where, and how food plants are grown and transported, as well as consider the amount consumed rela tive to that which is wasted. All food comes directly or indirectly from plants; roughly 10% of the known plant diversity is edible, and of these, 7000 species are cultivated for food (UNFAO, 2009, 2014). At the center of discussions about agriculture and
1Manuscript received 9 September 2014; revision accepted 26 September 2014. This Special Issue resulted in part from the combined symposium/ colloquium “Speaking of Food: Connecting Basic and Applied Plant Science” held at the 2013 Botany meetings in New Orleans, LA. Funding for the 2013 Speaking of Food symposium was generously provided by the Botanical Society of America’s Economic Botany Section, Genetics Section, and Systematics Section (with the American Society of Plant Taxonomists), and the Torrey Botanical Society. The editors are grateful to all of the speakers in the symposium, whose presentations inspired this issue and many other ongoing conversations about botany and its relationships to applied plant science as it relates to agriculture and agricultural impacts. The editors thank the many reviewers who thoughtfully reviewed earlier versions of the manuscripts in this issue. Finally, they thank Amy McPherson, Richard Hund, and the American Journal o f Botany staff for their support and editorial assistance. 5Author for correspondence (e-mail: [email protected]) doi:10.3732/ajb. 1400409
the future of food in a changing climate are the plants that we grow for food, fiber, and fuels (crops) and the science that is required to understand, improve, and conserve them. Since its inception 100 years ago, the American Journal o f Botany (AJB) has provided an important vehicle for the dis semination of plant science addressing a wide range of topics (structure, function, development, diversity, genetics, evolution, systematics) and levels of organization (molecular to ecosystem) (http://www.amjbot.org). An additional dimension of this char acteristic breadth is the inclusion of a breathtaking array of plant taxa, including studies of model plants (e.g., at least 76 papers on Arabidopsis have appeared in AJB, starting with Miksche and Brown, 1965), crops, ornamentals, and a wide diversity of wild species. AJB papers address fundamental questions in plant biology, using various techniques to advance basic under standing of how plants develop and function, how they interact with their abiotic and biotic environments, and how they are related to one another, among other things. These topics are crucial for understanding plants in general, but are also relevant for agriculture and the crop plants upon which we depend. This special issue of AJB, Speaking of Food: Connecting Basic and Applied Plant Science, aims to provide concrete examples of how different types of basic plant science, the types of scien tific studies commonly published in AJB, are relevant for the future of food. Indeed, AJB consistently publishes rigorous sci ence that pushes the boundaries of our understanding of bio logical processes in plants, critical determinants of plant survival and reproduction—papers about topics including pollination biology (Khorsand Rosa and Koptur, 2013), photosynthesis (Wang et al., 2008), water and nutrient transport (Suwa and Maherali, 2008), gene flow (Ellstrand, 2014), polyploidy
American Journal o f Botany 101(10): 1597-1600, 2014; http://www.amjbot.org/ © 2014 Botanical Society of America
1597
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(Barringer, 2007), and tropisms (Spalding, 2013). Research ad dressing these and other topics in crops and their wild relatives has figured prominently in the AJB in the past and present. Articles in the first issue of AJB dealt with floral and fruit development in ornamental plants (Gortner and Harris, 1914; White, 1914), and since then, hundreds of articles have been published on various crops and their wild relatives including Solarium (tomatoes and potatoes, at least 236 articles), com (116), wheat (97), soybean (40), rice (35), apples (35), and grapes (27), among many others. Building on this historical foundation, we challenged the contemporary AJB community to think about basic plant sci ence in the context of agriculture and the future of food. Con sideration of basic and applied plant research was inspired by Elizabeth Kellogg’s 2012 presidential address to the Botanical Society of America in Columbus, Ohio, when she pointed out that despite the importance of basic plant biology for agricul ture, it is rare that papers in AJB make this connection explicit. The issue presented here is part of a larger, more general dia logue happening in many academic communities and funding agencies: What is the connection between, and relationship of, basic and applied research? PASTEUR’S QUADRANT In an influential book in 1997, Donald Stokes introduced the idea of what he called “Pasteur’s quadrant” or use-inspired ba sic research. He argued that it is not possible or desirable to try to draw a bright line between “basic” and “applied” research. Instead, he suggested that it might be more productive to con sider two orthogonal axes, one reflecting the extent to which the research represents a quest for fundamental understanding and the other whether the research is inspired by immediate utility (Fig. 1). These axes define a set of quadrants, which Stokes
C onsideration of use? No
Yes
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Use-inspired basic research (Pasteur)
0
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0 Zi
a Fig. 1. Quadrant model of scientific research (modified from fig. 3-5 of Stokes, 1997), including exemplar practitioners. (Reprinted with per mission from Brookings Institution Press.)
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named after well-known scientists whose work exemplified his points. The upper left quadrant he called Bohr’s quadrant, for re search on basic principles (in this case, physics) that is under taken with no application in mind; it is driven by a quest for fundamental understanding but without regard for use. The lower right is Edison’s quadrant, which represents science under taken to find solutions to immediate problems but not particularly driven by a need to identify basic principles. Stokes did not name the lower left quadrant. The upper right quadrant, the focus of Stokes’s argument, is called Pasteur’s quadrant, or use-inspired basic research. This quadrant is the intersection of research undertaken with a desire for fundamental understanding of broad principles, but also with consideration of use. In the context of this Special Issue, we might interpret the term “fundamental understanding” to mean “broad biological significance” or principles that apply to a wide array of plant systems. In this issue, we aimed to illustrate that “basic” and “applied” are not two discrete categories, nor are they even extremes of a linear continuum. Rather, the terms can overlap such that re search that was once considered “basic” may ultimately be come critical for application to human-centered problems such as food production, and conversely, studies once classified as “applied” may illuminate general biological principles. Most of the plants discussed in this issue are crop plants or their close relatives; we argue that these fall firmly into Pasteur’s quadrant. While not included here for reasons of space, the same could be said of many studies of other systems, including noncrop models such as Arabidopsis or Brachypodium, and other wild plant species. OVERVIEW OF SPECIAL ISSUE Articles in this special issue span a broad range of topics showing the multitude of ways that research in “basic” plant science can contribute to and influence more “applied” research. Much of the research conducted in the plant sciences (and pub lished in AJB) approaches Pasteur’s quadrant, the use-inspired quadrant, when considered in the context of the need to feed a growing population on limited resources while still preserv ing plant diversity across the globe. Our hope is that this issue will inspire researchers working in a range of disciplines and systems to think about how their pursuit of basic scientific ad vances in plant science may have some relevance for sustainable agriculture. Several of the papers highlight the interactions between hu mans and plants, ranging from prehistoric to modem practices. The contributions of archaeobotany to unraveling the complex ities of crop origins, development, and (occasionally) abandon ment are detailed by Langlie et al. (2014). They showcase the importance of integrating information from multiple fields of study, demonstrate that great strides have been made in under standing domestication processes that are not linear, and pose intriguing (but rarely asked) questions about the culinary uses of the first domesticated plants. This theme is extended to the difficult case of species that were used (and perhaps incipiently domesticated), but which have been abandoned in recent times. Ross et al. (2014) demonstrate how patterns of human use can influence modern landscapes and how a variety of approaches are needed to distinguish between natural processes and past human intervention. Finally, the implications of current scien tific research for the future of food production and economic
October 2014]
G ross
et al .— I ntroduction
growth are explored by Reynolds et al. (2014), with an emphasis on the responsibility of plant scientists to consider how their research can and should be employed in the service of food production. All of these papers shed light on how humans have used plants through time and how human use of plants has in fluenced the environment up to the present. This research is a vital roadmap as we attempt to balance the needs of the human population with ecosystem function, a critical component of which is plant diversity. Understanding the diversity and evolutionary history of crops and their wild relatives is key to successfully managing existing domesticates and related species for crop improvement. These areas are addressed at several levels and in very different sys tems in this issue. Pearl and Burke (2014) document the genetic diversity of safflower (Carthamus tinctorius), a crop that is cur rently under-used but has the potential to be a major oilseed crop in the future, especially in dry climates. Sherman-Broyles et al. (2014) document 30 yr of research dedicated to under standing the origins, phylogenetic relationships, and genome structure of the perennial wild relatives of soybean, one of the world’s most important crops. Arbizu et al. (2014) explore the complex relationships among species and varieties of Daucus using 94 nuclear orthologs, and their findings shed light on both the structure of the genus and on how to efficiently use data in phylogenomics. Liston et al. (2014) survey the evolutionary, ecological, and genomic intricacies of domesticated and wild species of the genus Fragaria, showing how this group can link basic and applied research across disciplines. Ren et al. (2014) explore diversity in a neglected but critical group of crops and cultivated plants in southwestern China, documenting the pollination and breeding systems of these species for the first time in many cases. Taking a broader view that encompasses all angiosperms, Renny-Byfield and Wendel (2014) explore the important role of polyploidy in crop domestication and improvement. Domesticated species are never free from their weedy com panions, and an enormous amount of time, money, and effort is directed toward controlling or eradicating weeds; agriculture and food production cannot be discussed without reference to these influential plants. Waselkov and Olsen (2014) survey the population genetics of waterhemp (Amaranthus tuberculatus), a weed that is native to the Midwest and that has become in creasingly problematic for US farmers since the 1950s. This work shows that a genetic lineage from the western United States has expanded to the east along with modem farming techniques and is the source of most weedy populations. Thurber et al. (2014) explore flowering time in weedy rice (conspecific with domesticated rice, Oryza sativa), one of the traits that makes weedy rice an effective weed. Their work shows that there was rapid divergence in weedy rice flowering time, such that some weeds flower earlier and some later than domesticated rice. Nonetheless, flowering times still overlap (with the potential for crop-weed hybridization), and there is allele sharing between the crop and the weed at major loci controlling flowering. Other papers address the question of how crops and their wild relatives respond to natural and artificial selective pres sures. Henry et al. (2014) approach this through genetic charac terization of variable flowering time in wild sunflower. They find multiple origins of short-day induced flowering and fairly simple genetic control of flowering, suggesting that this varia tion could be easily introgressed into domesticated sunflower. Doust et al. (2014) document the basis of the economically criti cal domestication trait of loss-of-shattering across the grasses.
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The broad survey shows that while nonshattering is derived from a variety of shattering phenotypes, there is support for a common genetic pathway, and they propose future directions to test this possibility rigorously. Finally, several papers return to the question of human and plant interactions, documenting the genetic implications of do mestication and improvement for crop species, and asking how this information can be applied to the future of crop develop ment. Gross et al. (2014) focus on the impact of hundreds of years of crop breeding on the genetic diversity of domesticated apple in a comparison of annual and perennial crops. Apples show almost no loss of diversity through time, although the genetic diversity of the most commonly grown cultivars is much lower. McClure et al. (2014) deal with the related prob lem of perennial crop breeding. Despite the vast genetic diver sity available in many perennial crops, most of it is not used due to the challenges associated with generation length in perennial species. They show that genomic tools may provide a way out of the constantly narrowing genetic diversity of elite perennial cultivars. Warschefsky et al. (2014) explore the rich resources available from systematic hybridization between crops and their wild relatives. This variation may sometimes be challeng ing to access, but it could be a critical resource for responding to the increasing challenges in food production. DeHaan and Van Tassel (2014) depart from the improvement of existing crops and seek instead to apply existing information about crop domestication and evolution to the domestication of wholly new perennial grain crops.
LOOKING FORWARD Developing sustainable agriculture for a growing population is a primary challenge facing society. Applied science aimed at crop improvement and the enhancement of particular agro nomic inputs is critical to the solution. Here, we argue that basic plant research focused on expanding general understanding of biological phenomena and processes in plants is important for agriculture as well. This work can be carried out in crops and their wild and weedy relatives, model plants, or any plant spe cies that offers an opportunity to address important questions about the biology of plants. We believe that many facets of con temporary plant science have a place in Pasteur’s quadrant, where use-inspired basic research pushes the boundaries of un derstanding while simultaneously contributing valuable infor mation relevant for the future of food. LITERATURE CITED A rbizu, C., H. R uess, D. S enalik, P. W. S imon, and D. M. S pooner. 2014.
Phylogenomics of the carrot genus (Daucus, Apiaceae). American Journal o f Botany 101: 1666-1685. B arringer, B. C. 2007. Polyploidy and self-fertilization in flowering plants. American Journal of Botany 94: 1527-1533. D eH aan, L. R., and D. L. V an T assel. 2014. Useful insights from evo lutionary biology for developing perennial grain crops. American Journal o f Botany 101: 1801-1819. D oust, A. N., M. M auro-H errera, A. D. F rancis, and L. C. S hand. 2014. Morphological diversity and genetic regulation of inflores cence abscission zones in grasses. American Journal of Botany 101: 1759-1769. E llstrand, N. C. 2014. Is gene flow the most important evolutionary force in plants? American Journal of Botany 101: 737-753.
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A merican J ournal of B otany
and J. A. H arris. 1914. On axial abscission in Impatiens sultani as the result of traumatic stimuli. American Journal o f Botany 1: 48-50. G ross, B. L., A. D. H enk, C. M. Richards, G. F azio, and G. M. V olk. 2014. Genetic diversity in Malus xdomestica (Rosaceae) through time in re sponse to domestication. American Journal of Botany 101: 1770-1779. H enry, L. R, R. H. B. W atson, and B. K. B lackman. 2014. Transitions in photoperiodic flowering are common and involve few loci in wild sunflowers (Helianthus; Asteraceae). American Journal o f Botany 101: 1748-1758. K ellogg, E. A. 2012. Botany 2012 presidential address: “Speaking of food”. Plant Science Bulletin 58(3): 78-81. Khorsand R osa, R. K., and S. K optur. 2013. New findings on the pol lination biology of Mauritiaflexuosa (Arecaceae) in Roraima, Brazil: Linking dioecy, wind, and habitat. American Journal of Botany 100: 613-621. L anglie, B. S., N. G. M ueller, R. N. S pengler, and G. J. F ritz. 2014. Agricultural origins from the ground up: Archaeological approaches to plant domestication. American Journal o f Botany 101: 1601-1617. L iston, A., R. C ronn, and T.-L. A shman. 2014. Fragaria: A genus with deep historical roots and ripe for evolutionary and ecological insights. American Journal o f Botany 101: 1686-1699. M cC lure, K. A., J. S awler, K. M. G ardner, D. M oney, and S. M yles. 2014. Genomics: A potential panacea for the perennial problem. American Journal o f Botany 101: 1780-1790. M iksche, J. P., and J. A. M. B rown. 1965. Development of vegetative and floral meristems of Arabidopsis thaliana. American Journal of Botany 52: 533-537. P earl, S. A., and J. M. B urke. 2014. Genetic diversity in Carthamus tinctorius (Asteraceae; safflower), an underutilized oilseed crop. American Journal o f Botany 101: 1640-1650. Ren, Z.-X., H. W ang, P. B ernhardt, and D.-Z. Li. 2014. Insect pol lination and self-incompatibility in edible and/or medicinal crops in southwestern China, a global hotspot of biodiversity. American Journal of Botany 101: 1700-1710. R enny-B yfield, S., and J. F. W endel. 2014. Doubling down on ge nomes: Polyploidy and crop plants. American Journal of Botany 101: 1711-1725. R eynolds, H. L., A. A. S mith, and J. R. F armer. 2014. Think globally, research locally: Paradigms and place in agroecological research. American Journal o f Botany 101: 1631-1639.
Ross, N. J., M. H. H. Stevens, A. W. R upiper, I. H arkreader, and L. A. L eben. 2014. The ecological side of an ethnobotanical coin: Legacies in historically managed trees. American Journal o f Botany 101: 1618-1630. S herman-B royles, S., A. B ombarely, A. F. P owell, J. L. D oyle, A. N. E gan, J. E. C oate, and J. J. D oyle. 2014. The wild side of a ma jor crop: Soybean’s perennial cousins from Down Under. American Journal o f Botany 101: 1651-1665. Spalding, E. P. 2013. Diverting the downhill flow of auxin to steer growth during tropisms. American Journal o f Botany 100: 203-214. S tokes, D. E. 1997. Pasteur’s quadrant—Basic science and technological innovation. Brookings Institution Press, Washington D.C., USA. S uwa, T., and H. M aherali. 2008. Influence of nutrient availability on the mechanisms of tolerance to herbivory in an annual grass, Avena barbata (Poaceae). American Journal o f Botany 95: 434-440. T hurber, C. S., M. R eagon, K. M. O lsen, Y. J ia , and A. L. C aicedo. 2014. The evolution of flowering strategies in US weedy rice. American Journal o f Botany 101: 1737-1747. UNFAO [U nited N ational F ood and A griculture O rganization], 2009. High level expert forum—How to feed the world in 2050. Economic and Social Development Department, Food and Agriculture Organization, Rome, Italy. UNFAO [United N ations F ood
and
A griculture O rganization]. 2014.
Biodiversity, components, plants. Available at http://www.fao.org/ biodiversity/components/plants/en. Wang, D., S. A. H eckathorn, D. B arua, P. J oshi, E. W. H amilton, and J. J. L acroix. 2008. Effects of elevated C 0 2 on the tolerance of pho tosynthesis to acute heat stress in C3, C4, and CAM species. American Journal of Botany 95: 165-176. W arschefsky, E., R. V. P enmetsa, D. R. C ook, and E. J. B. von W ettberg. 2014. Back to the wilds: Tapping evolutionary adaptations for resil ient crops through systematic hybridization with crop wild relatives. American Journal o f Botany 101: 1791-1800. W aselkov, K. E., and K. M. O lsen. 2014. Population genetics and origin of the native North American agricultural weed waterhemp (Amaranthus tuberculatus; Amaranthaceae). American Journal o f Botany 101: 1726-1736. W hite, O. 1914. Studies of teratological phenomena in their relation to evolution and the problems of heredity. I. A study of certain floral ab normalities in Nicotiana and their bearing on theories of dominance. American Journal o f Botany 1: 23-36.
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American Journal of Botany 101(10): 1597-1600,2014.
C e le b r a t in g lO O y e a r s ' ^ 11 9 1 4 - 2 0 1 4
INTRODUCTION
S peaking of food: C onnecting basic and applied plant science1 B riana
L. G ross2, E lizabeth A.
K ellogg 3,
and
A llison
J. M iller4’5
departm ent of Biology, University of Minnesota Duluth, 207 Swenson Science Building, 1035 Kirby Drive, Duluth Minnesota 55812 USA; 3Donald Danforth Plant Science Center, 975 North Warson Road, St. Louis, Missouri 63132 USA; and departm ent of Biology, Saint Louis University, 3507 Laclede Avenue, St. Louis, Missouri 63103-2010 USA The Food and Agriculture Organization (FAO) predicts that food production must rise 70% over the next 40 years to meet the demands of a growing population that is expected to reach nine billion by the year 2050. Many facets of basic plant science promoted by the Botanical Society of America are important for agriculture; however, more explicit connections are needed to bridge the gap between basic and applied plant research. This special issue, Speaking of Food: Connecting Basic and Applied Plant Science, was conceived to showcase productive overlaps of basic and applied research to address the challenges posed by feeding billions of people and to stimulate more research, fresh connections, and new paradigms. Contributions to this special issue thus illustrate some interactive areas of study in plant science— historical and modem plant-human interaction, crop and weed origins and evolution, and the effects of natural and artificial selection on crops and their wild relatives. These papers provide examples of how research integrating the basic and applied aspects of plant science benefits the pursuit of knowledge and the translation of that knowledge into actions toward sustainable production of crops and conservation of diversity in a changing climate.
Key words: agriculture; agroecological research; applied science; basic science; botany; crop domestication; crop evolution; food production; Pasteur’s quadrant. The Food and Agriculture Organization of the United Na tions predicts that food production must rise by 70% over the next 40 yr to meet the demands of a growing population ex pected to reach nine billion by the year 2050 (UNFAO, 2009). The solution to this challenge will be multifaceted and certainly involve changes in what, where, and how food plants are grown and transported, as well as consider the amount consumed rela tive to that which is wasted. All food comes directly or indirectly from plants; roughly 10% of the known plant diversity is edible, and of these, 7000 species are cultivated for food (UNFAO, 2009, 2014). At the center of discussions about agriculture and
1Manuscript received 9 September 2014; revision accepted 26 September 2014. This Special Issue resulted in part from the combined symposium/ colloquium “Speaking of Food: Connecting Basic and Applied Plant Science” held at the 2013 Botany meetings in New Orleans, LA. Funding for the 2013 Speaking of Food symposium was generously provided by the Botanical Society of America’s Economic Botany Section, Genetics Section, and Systematics Section (with the American Society of Plant Taxonomists), and the Torrey Botanical Society. The editors are grateful to all of the speakers in the symposium, whose presentations inspired this issue and many other ongoing conversations about botany and its relationships to applied plant science as it relates to agriculture and agricultural impacts. The editors thank the many reviewers who thoughtfully reviewed earlier versions of the manuscripts in this issue. Finally, they thank Amy McPherson, Richard Hund, and the American Journal o f Botany staff for their support and editorial assistance. 5Author for correspondence (e-mail: [email protected]) doi:10.3732/ajb. 1400409
the future of food in a changing climate are the plants that we grow for food, fiber, and fuels (crops) and the science that is required to understand, improve, and conserve them. Since its inception 100 years ago, the American Journal o f Botany (AJB) has provided an important vehicle for the dis semination of plant science addressing a wide range of topics (structure, function, development, diversity, genetics, evolution, systematics) and levels of organization (molecular to ecosystem) (http://www.amjbot.org). An additional dimension of this char acteristic breadth is the inclusion of a breathtaking array of plant taxa, including studies of model plants (e.g., at least 76 papers on Arabidopsis have appeared in AJB, starting with Miksche and Brown, 1965), crops, ornamentals, and a wide diversity of wild species. AJB papers address fundamental questions in plant biology, using various techniques to advance basic under standing of how plants develop and function, how they interact with their abiotic and biotic environments, and how they are related to one another, among other things. These topics are crucial for understanding plants in general, but are also relevant for agriculture and the crop plants upon which we depend. This special issue of AJB, Speaking of Food: Connecting Basic and Applied Plant Science, aims to provide concrete examples of how different types of basic plant science, the types of scien tific studies commonly published in AJB, are relevant for the future of food. Indeed, AJB consistently publishes rigorous sci ence that pushes the boundaries of our understanding of bio logical processes in plants, critical determinants of plant survival and reproduction—papers about topics including pollination biology (Khorsand Rosa and Koptur, 2013), photosynthesis (Wang et al., 2008), water and nutrient transport (Suwa and Maherali, 2008), gene flow (Ellstrand, 2014), polyploidy
American Journal o f Botany 101(10): 1597-1600, 2014; http://www.amjbot.org/ © 2014 Botanical Society of America
1597
1598
A merican J ournal
(Barringer, 2007), and tropisms (Spalding, 2013). Research ad dressing these and other topics in crops and their wild relatives has figured prominently in the AJB in the past and present. Articles in the first issue of AJB dealt with floral and fruit development in ornamental plants (Gortner and Harris, 1914; White, 1914), and since then, hundreds of articles have been published on various crops and their wild relatives including Solarium (tomatoes and potatoes, at least 236 articles), com (116), wheat (97), soybean (40), rice (35), apples (35), and grapes (27), among many others. Building on this historical foundation, we challenged the contemporary AJB community to think about basic plant sci ence in the context of agriculture and the future of food. Con sideration of basic and applied plant research was inspired by Elizabeth Kellogg’s 2012 presidential address to the Botanical Society of America in Columbus, Ohio, when she pointed out that despite the importance of basic plant biology for agricul ture, it is rare that papers in AJB make this connection explicit. The issue presented here is part of a larger, more general dia logue happening in many academic communities and funding agencies: What is the connection between, and relationship of, basic and applied research? PASTEUR’S QUADRANT In an influential book in 1997, Donald Stokes introduced the idea of what he called “Pasteur’s quadrant” or use-inspired ba sic research. He argued that it is not possible or desirable to try to draw a bright line between “basic” and “applied” research. Instead, he suggested that it might be more productive to con sider two orthogonal axes, one reflecting the extent to which the research represents a quest for fundamental understanding and the other whether the research is inspired by immediate utility (Fig. 1). These axes define a set of quadrants, which Stokes
C onsideration of use? No
Yes
05
c
'tj
c
B
« 0 CD >0
T3
c zs
Pure basic research (Bohr)
Use-inspired basic research (Pasteur)
0
c 0 E 0
■O
c 0
w
o z
Pure applied research (Edison)
0 Zi
a Fig. 1. Quadrant model of scientific research (modified from fig. 3-5 of Stokes, 1997), including exemplar practitioners. (Reprinted with per mission from Brookings Institution Press.)
of
[Vol. 101
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named after well-known scientists whose work exemplified his points. The upper left quadrant he called Bohr’s quadrant, for re search on basic principles (in this case, physics) that is under taken with no application in mind; it is driven by a quest for fundamental understanding but without regard for use. The lower right is Edison’s quadrant, which represents science under taken to find solutions to immediate problems but not particularly driven by a need to identify basic principles. Stokes did not name the lower left quadrant. The upper right quadrant, the focus of Stokes’s argument, is called Pasteur’s quadrant, or use-inspired basic research. This quadrant is the intersection of research undertaken with a desire for fundamental understanding of broad principles, but also with consideration of use. In the context of this Special Issue, we might interpret the term “fundamental understanding” to mean “broad biological significance” or principles that apply to a wide array of plant systems. In this issue, we aimed to illustrate that “basic” and “applied” are not two discrete categories, nor are they even extremes of a linear continuum. Rather, the terms can overlap such that re search that was once considered “basic” may ultimately be come critical for application to human-centered problems such as food production, and conversely, studies once classified as “applied” may illuminate general biological principles. Most of the plants discussed in this issue are crop plants or their close relatives; we argue that these fall firmly into Pasteur’s quadrant. While not included here for reasons of space, the same could be said of many studies of other systems, including noncrop models such as Arabidopsis or Brachypodium, and other wild plant species. OVERVIEW OF SPECIAL ISSUE Articles in this special issue span a broad range of topics showing the multitude of ways that research in “basic” plant science can contribute to and influence more “applied” research. Much of the research conducted in the plant sciences (and pub lished in AJB) approaches Pasteur’s quadrant, the use-inspired quadrant, when considered in the context of the need to feed a growing population on limited resources while still preserv ing plant diversity across the globe. Our hope is that this issue will inspire researchers working in a range of disciplines and systems to think about how their pursuit of basic scientific ad vances in plant science may have some relevance for sustainable agriculture. Several of the papers highlight the interactions between hu mans and plants, ranging from prehistoric to modem practices. The contributions of archaeobotany to unraveling the complex ities of crop origins, development, and (occasionally) abandon ment are detailed by Langlie et al. (2014). They showcase the importance of integrating information from multiple fields of study, demonstrate that great strides have been made in under standing domestication processes that are not linear, and pose intriguing (but rarely asked) questions about the culinary uses of the first domesticated plants. This theme is extended to the difficult case of species that were used (and perhaps incipiently domesticated), but which have been abandoned in recent times. Ross et al. (2014) demonstrate how patterns of human use can influence modern landscapes and how a variety of approaches are needed to distinguish between natural processes and past human intervention. Finally, the implications of current scien tific research for the future of food production and economic
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growth are explored by Reynolds et al. (2014), with an emphasis on the responsibility of plant scientists to consider how their research can and should be employed in the service of food production. All of these papers shed light on how humans have used plants through time and how human use of plants has in fluenced the environment up to the present. This research is a vital roadmap as we attempt to balance the needs of the human population with ecosystem function, a critical component of which is plant diversity. Understanding the diversity and evolutionary history of crops and their wild relatives is key to successfully managing existing domesticates and related species for crop improvement. These areas are addressed at several levels and in very different sys tems in this issue. Pearl and Burke (2014) document the genetic diversity of safflower (Carthamus tinctorius), a crop that is cur rently under-used but has the potential to be a major oilseed crop in the future, especially in dry climates. Sherman-Broyles et al. (2014) document 30 yr of research dedicated to under standing the origins, phylogenetic relationships, and genome structure of the perennial wild relatives of soybean, one of the world’s most important crops. Arbizu et al. (2014) explore the complex relationships among species and varieties of Daucus using 94 nuclear orthologs, and their findings shed light on both the structure of the genus and on how to efficiently use data in phylogenomics. Liston et al. (2014) survey the evolutionary, ecological, and genomic intricacies of domesticated and wild species of the genus Fragaria, showing how this group can link basic and applied research across disciplines. Ren et al. (2014) explore diversity in a neglected but critical group of crops and cultivated plants in southwestern China, documenting the pollination and breeding systems of these species for the first time in many cases. Taking a broader view that encompasses all angiosperms, Renny-Byfield and Wendel (2014) explore the important role of polyploidy in crop domestication and improvement. Domesticated species are never free from their weedy com panions, and an enormous amount of time, money, and effort is directed toward controlling or eradicating weeds; agriculture and food production cannot be discussed without reference to these influential plants. Waselkov and Olsen (2014) survey the population genetics of waterhemp (Amaranthus tuberculatus), a weed that is native to the Midwest and that has become in creasingly problematic for US farmers since the 1950s. This work shows that a genetic lineage from the western United States has expanded to the east along with modem farming techniques and is the source of most weedy populations. Thurber et al. (2014) explore flowering time in weedy rice (conspecific with domesticated rice, Oryza sativa), one of the traits that makes weedy rice an effective weed. Their work shows that there was rapid divergence in weedy rice flowering time, such that some weeds flower earlier and some later than domesticated rice. Nonetheless, flowering times still overlap (with the potential for crop-weed hybridization), and there is allele sharing between the crop and the weed at major loci controlling flowering. Other papers address the question of how crops and their wild relatives respond to natural and artificial selective pres sures. Henry et al. (2014) approach this through genetic charac terization of variable flowering time in wild sunflower. They find multiple origins of short-day induced flowering and fairly simple genetic control of flowering, suggesting that this varia tion could be easily introgressed into domesticated sunflower. Doust et al. (2014) document the basis of the economically criti cal domestication trait of loss-of-shattering across the grasses.
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The broad survey shows that while nonshattering is derived from a variety of shattering phenotypes, there is support for a common genetic pathway, and they propose future directions to test this possibility rigorously. Finally, several papers return to the question of human and plant interactions, documenting the genetic implications of do mestication and improvement for crop species, and asking how this information can be applied to the future of crop develop ment. Gross et al. (2014) focus on the impact of hundreds of years of crop breeding on the genetic diversity of domesticated apple in a comparison of annual and perennial crops. Apples show almost no loss of diversity through time, although the genetic diversity of the most commonly grown cultivars is much lower. McClure et al. (2014) deal with the related prob lem of perennial crop breeding. Despite the vast genetic diver sity available in many perennial crops, most of it is not used due to the challenges associated with generation length in perennial species. They show that genomic tools may provide a way out of the constantly narrowing genetic diversity of elite perennial cultivars. Warschefsky et al. (2014) explore the rich resources available from systematic hybridization between crops and their wild relatives. This variation may sometimes be challeng ing to access, but it could be a critical resource for responding to the increasing challenges in food production. DeHaan and Van Tassel (2014) depart from the improvement of existing crops and seek instead to apply existing information about crop domestication and evolution to the domestication of wholly new perennial grain crops.
LOOKING FORWARD Developing sustainable agriculture for a growing population is a primary challenge facing society. Applied science aimed at crop improvement and the enhancement of particular agro nomic inputs is critical to the solution. Here, we argue that basic plant research focused on expanding general understanding of biological phenomena and processes in plants is important for agriculture as well. This work can be carried out in crops and their wild and weedy relatives, model plants, or any plant spe cies that offers an opportunity to address important questions about the biology of plants. We believe that many facets of con temporary plant science have a place in Pasteur’s quadrant, where use-inspired basic research pushes the boundaries of un derstanding while simultaneously contributing valuable infor mation relevant for the future of food. LITERATURE CITED A rbizu, C., H. R uess, D. S enalik, P. W. S imon, and D. M. S pooner. 2014.
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